Aedes albopictus colonies from different geographic origins differ in their sleep and activity levels but not in the time of peak activity.

Mosquitoes occupy a wide range of habitats where they experience various environmental conditions. The ability of some species, such as the tiger mosquito, Aedes albopictus, to adapt to local conditions certainly contributes to their invasive success. Among traits that remain to be examined, mosquitoes' ability to time their activity with that of the local host population has been suggested to be of significant epidemiological importance. However, whether different populations display heritable differences in their chronotype has not been examined. Here, we compared laboratory strains originating from 8 populations from 3 continents, monitored their spontaneous locomotor activity patterns, and analyzed their sleep-like states. Overall, all strains showed conserved diurnal activity concentrated in the hours preceding the crepuscule. Similarly, they all showed increased sleep levels during the morning and night hours. However, we observed strain-specific differences in the activity levels at each phase of the day. We also observed differences in the fraction of time that each strain spends in a sleep-like state, explained by variations in the sleep architecture across strains. Human population density and the latitude of the site of geographic origin of the tested strain showed significant effects on sleep and activity patterns. Altogether, these results suggest that Ae. albopictus mosquitoes adapt to local environmental conditions via heritable adaptations of their chronotype.

Visual threats reduce blood-feeding and trigger escape responses in Aedes aegypti mosquitoes.

The diurnal mosquitoes Aedes aegypti are vectors of several arboviruses, including dengue, yellow fever, and Zika viruses. To find a host to feed on, they rely on the sophisticated integration of olfactory, visual, thermal, and gustatory cues emitted by the hosts. If detected by their target, this latter may display defensive behaviors that mosquitoes need to be able to detect and escape in order to survive. In humans, a typical response is a swat of the hand, which generates both mechanical and visual perturbations aimed at a mosquito. Here, we used programmable visual displays to generate expanding objects sharing characteristics with the visual component of an approaching hand and quantified the behavioral response of female mosquitoes. Results show that Ae. aegypti is capable of using visual information to decide whether to feed on an artificial host mimic. Stimulations delivered in a LED flight arena further reveal that landed Ae. aegypti females display a stereotypical escape strategy by taking off at an angle that is a function of the direction of stimulus introduction. Altogether, this study demonstrates that mosquitoes landed on a host mimic can use isolated visual cues to detect and avoid a potential threat.

Species-Specificity in Thermopreference and CO2-Gated Heat-Seeking in Culex Mosquitoes.

Combining thermopreference (Tp) and CO2-gated heat-seeking assays, we studied the thermal preferendum and response to thermal cues in three Culex mosquito species exhibiting differences in native habitat and host preference (e.g., biting cold and/or warm-blooded animals). Results show that these species differ in both Tp and heat-seeking behavior. In particular, we found that Culex territans, which feed primarily on cold-blood hosts, did not respond to heat during heat-seeking assays, regardless of the CO2 concentration, but exhibited an intermediate Tp during resting. In contrast, Cx. quinquefasciatus, which feeds on warm blooded hosts, sought the coolest locations on a thermal gradient and responded only moderately to thermal stimuli when paired with CO2 at higher concentrations. The third species, Cx. tarsalis, which has been shown to feed on a wide range of hosts, responded to heat when paired with high CO2 levels and exhibited a high Tp. This study provides the first insights into the role of heat and CO2 in the host seeking behavior of three disease vectors in the Culex genus and highlights differences in preferred resting temperatures.

Modeling the effects of Aedes aegypti's larval environment on adult body mass at emergence.

Mosquitoes vector harmful pathogens that infect millions of people every year, and developing approaches to effectively control mosquitoes is a topic of great interest. However, the success of many control measures is highly dependent upon ecological, physiological, and life history traits of mosquito species. The behavior of mosquitoes and their potential to vector pathogens can also be impacted by these traits. One trait of interest is mosquito body mass, which depends upon many factors associated with the environment in which juvenile mosquitoes develop. Our experiments examined the impact of larval density on the body mass of Aedes aegypti mosquitoes, which are important vectors of dengue, Zika, yellow fever, and other pathogens. To investigate the interactions between the larval environment and mosquito body mass, we built a discrete time mathematical model that incorporates body mass, larval density, and food availability and fit the model to our experimental data. We considered three categories of model complexity informed by data, and selected the best model within each category using Akaike's Information Criterion. We found that the larval environment is an important determinant of the body mass of mosquitoes upon emergence. Furthermore, we found that larval density has greater impact on body mass of adults at emergence than on development time, and that inclusion of density dependence in the survival of female aquatic stages in models is important. We discuss the implications of our results for the control of Aedes mosquitoes and on their potential to spread disease.

Antipredatory Responses of Mosquito Pupae to Non-Lethal Predation Threat-Behavioral Plasticity Across Life-History Stages.

Antipredatory behavioral responses tend to be energetically expensive, and prey species thus need to resolve trade-offs between these behaviors and other activities such as foraging and mating. While these trade-offs have been well-studied across taxa, less is known about how costs and benefits vary in different life-history contexts, and associated consequences. To address this question, we compared responses of the yellow fever mosquito (Aedes aegypti [Diptera: Culicidae]) to predation threat from guppy (Poecilia reticulata [Cyprinodontiformes: Poeciliidae]) across two life-history stages-larvae (data from previous study) and pupae (from this study). Pupae are motile but do not feed and are comparable to larvae in terms of behavior. To understand how physiological costs affect the threat sensitivity of pupae, we used sex (with size as a covariate) as a proxy for stored energy reserves, and quantified movement and space use patterns of male (small-sized) and female (large-sized) pupae when exposed to predation threat. We found that pupae did not alter movement when exposed to predator cues but instead altered spatial use by spending more time at the bottom of the water column. We found no effect of pupa sex (or size) on the behavioral responses we measured. We conclude that pupa behavior, both antipredatory and otherwise, is primarily targeted at minimizing energy expenditure, as compared with larval behavior, which appears to balance energy expenditure between the opposing pressures of foraging and of avoiding predation. We suggest that antipredatory defenses in metamorphosing prey are modulated by varying energetic trade-offs associated with different life-history stages.

Linking Mosquito Ecology, Traits, Behavior, and Disease Transmission.

Mosquitoes are considered to be the deadliest animals on Earth because the diseases they transmit claim at least a million human lives every year globally. Here, we discuss the scales at which the effects of ecological factors cascade to influence epidemiologically relevant behaviors of adult mosquitoes. In particular, we focused our review on the environmental conditions (coarse-scale variables) that shape the life-history traits of larvae and adult mosquitoes (fine-scale traits), and how these factors and their association, in turn, modulate adult behaviors to influence mosquito-borne disease transmission. Finally, we explore the integration of physical, physiological, and behavioral information into predictive models with epidemiological applications.

How Do Trait-Mediated Non-lethal Effects of Predation Affect Population-Level Performance of Mosquitoes?

Non-lethal, trait-mediated effects of predation impact prey behavior and life-history traits. Studying how these effects in turn influence prey demography is crucial to understand prey life-history evolution. Mosquitoes are important vectors that claim several million lives every year worldwide by transmitting a range of pathogens. Several ecological factors affect life-history traits of both larval and adult mosquitoes, creating effects that cascade to population-level consequences. Few studies have comprehensively explored the non-lethal effects of predation and its interactions with resources and competition on larval, adult, and population traits of mosquitoes. Understanding these interactions is important because the effects of predation are hypothesized to rescue prey populations from the effects of density-dependence resulting from larval competition. Aedes aegypti larvae reared at two different larval densities and subjected to three non-lethal predator treatments were monitored for survival, development time, and adult size through the larval stages to adult eclosion, and adult females were monitored for survival and reproduction through their first gonotrophic cycle. Intraspecific competition increased larval development time, yielded small-bodied adults, and reduced fecundity in individuals exposed to predatory chemical cues as larvae. Exposure to cues from a living predator affected both body size and latency to blood feed in females. Analysis of life-table traits revealed significant effects of competition on net reproductive rate (R 0) of mosquitoes. The interaction between competition and predator treatments significantly affected the cohort rate of increase (r) and the index of performance (r'). The index of performance, which estimates rate of population change based on the size-fecundity relationship, was significantly and positively correlated with r, but overestimated r slightly. Lack of significant effect of predator treatments and larval density on cohort generation time (T c) further suggests that the observed effects of treatments on r and r' were largely a consequence of the effects on R 0. Also, the significant effects of treatment combinations on larval development time, adult body size and fecundity were ultimately manifested as effects on life-table traits estimated from adult survival and reproduction.

Context-dependent interactive effects of non-lethal predation on larvae impact adult longevity and body composition.

Predation impacts development, behavior and morphology of prey species thereby shaping their abundances, distribution and community structure. Non-lethal threat of predation, specifically, can have a strong influence on prey lifehistory characteristics. While investigations often focus on the impact of predation threat on prey in isolation, tests of its interactive effects with food availability and resource competition on prey survival and fitness can improve understanding of costs, benefits and trade-offs of anti-predator strategies. This study, involving Aedes aegypti mosquitoes as a model organism, investigates both simple and interactive effects of predation threat during the larval stage on survival, size at and time to maturity, stored teneral reserves of glycogen, protein and lipid in adults, and adult longevity. Our results show that development times of mosquito larvae were increased (by 14.84% in males and by 97.63% in females), and size of eclosing adults decreased (by 62.30% in males and by 58.33% in females) when exposed to lowered nutrition and elevated intraspecific competition, but that predation had no detectable effect on these simple traits. Teneral reserves of glycogen, protein and lipid and adult longevity were positively correlated with adult body size. Non-lethal predation threat had significant interactive effects with nutrition and larval competition on teneral reserves in males and adult longevity in males and females. The sexes responded differently to conditions encountered as larvae, with the larval environment affecting development and adult characteristics more acutely for females than for males. The outcome of this study shows how threat of predation on juveniles can have long-lasting effects on adults that are likely to impact mosquito population dynamics and that may impact disease transmission.