Differential survival of Aedes aegypti and Aedes albopictus (Diptera: Culicidae) larvae exposed to low temperatures in Taiwan

Aedes aegypti (L.) and Aedes albopictus (Skuse) differ in their distribution in Taiwan. The former species is distributed in the south of Taiwan, whereas the latter is found throughout the island. One possible explanation proposes that low temperatures in the winter prevent the expansion of Ae. aegypti. Hence, the impact of low temperatures on immatures of both species was studied in the laboratory and in the field. Our study showed that, under most conditions, Ae. aegypti were more sensitive to low temperatures than Ae. albopictus both in the laboratory and in the field. The survival rates at 10 degrees C for the first and fourth instars of Ae. albopictus were significantly better than those of Ae. aegypti. At 2.5 and 5.0 degrees C, the first instars of Ae. albopictus survived better than the same stadium of Ae. aegypti, but the fourth instars of Ae. aegypti survived better. Short exposures to low temperatures did not affect the acclimatization of Ae. aegypti immatures but longer exposures did increase the physiological adaptation to low temperatures. For Ae. albopictus, exposure to low temperatures increases the acclimatization of this species. In field experiments, Ae. aegypti larvae had a significantly higher mortality than Ae. albopictus during exposures to cold fronts in the 2004 winter. We conclude that low temperatures in northern and central Taiwan have a negative impact on the distribution of Ae. aegypti, but this factor alone is not sufficient to prevent this species from occupying the rest of Taiwan.     

Intraguild predation among larval treehole mosquitoes, Aedes albopictus, Ae. aegypti, and Ae. triseriatus (Diptera: Culicidae), in laboratory microcosms.

We compared the tendency for 4th-instar larvae to prey on newly hatched larvae, and the vulnerability of those 1st instars to such predation for Aedes triseriatus (Say), Ae. aegypti (L.), and Ae. albopictus (Skuse), all container-breeding mosquitoes. The latter 2 species were introduced to North America and are now sympatric with Ae. triseriatus, a native species in eastern North America. The experiment also enabled the assessment of species-specific influences of food supplements and spatial heterogeneity on predatory behavior. Ae. triseriatus was substantially more predatory and less susceptible to attack than the other 2 species. These differences were amplified in food-deprived and spatially simple conditions, indicating that Ae. triseriatus predatory behavior may have important retarding effects on the colonization of occupied treehole habitats by Ae. albopictus. Ae. aegypti and Ae. albopictus were similar in imposing little (Ae. aegypti) or almost no (Ae. albopictus) predation on 1st instars and in being susceptible to predation by Ae. triseriatus. The general lack of species-specific differences between Ae. aegypti and Ae. albopictus indicates that interspecific predation is not a likely explanation for the rapid displacement of Ae. aegypti by Ae. albopictus in domestic containers in the southeastern United States.     

Laboratory study of competition between United States strains of Aedes albopictus and Aedes aegypti (Diptera: Culicidae)

Mosquito abatement workers in Houston, Tex., and New Orleans, La., have observed that the recent introduction of Aedes albopictus (Skuse) has been accompanied by a decline and virtual disappearance of Ae. aegypti (L.). This suggests competitive displacement but contradicts the direction of displacement observed in native habitats of Ae. albopictus. The paper reports a series of laboratory studies on competition between U.S. strains of the two species. Larval survivorship, size, and development time were monitored in increasing densities of pure and mixed cultures of the two species under limited and optimal diets. Oviposition preferences were examined by offering gravid females cups containing first or fourth instars of either species. Ethological isolation was studied by simultaneously offering virgin females of one species to males of both species and monitoring courtship behavior as well as rates of oviposition, percentage of hatch, and embryonation. The results of the larval competition studies suggest that interspecific competition increased development time only slightly and had no effect on survival of Ae. albopictus. Competition increased survival in Ae. aegypti. Size of both species was influenced by interspecific competition, but the magnitude and direction of the effect was inconsistent between strains and diets such that neither species emerged consistently larger. The results of the oviposition study indicated a higher fecundity in Ae. aegypti but otherwise demonstrated that both species oviposit without regard to the presence, age, or species of larvae in the oviposition container. The ethological isolation tests demonstrated that Ae. aegypti males were more sexually aggressive than males or Ae. albopictus. However, Ae. albopitus females placed with increasing densities of Ae. aegypti males demonstrated no decrease in rates of oviposition, and eggs laid showed equal rates of hatch and embryonation. In general, the results do not suggest that the U.S Ae. albopictus population is inherently more competitive in the laboratory than Ae. aegypti. Other reasons for the observed decline in Ae. aegypti in the United States are discussed.     

Interspecific competition among Aedes aegypti, Ae. albopictus, and Ae. triseriatus (Diptera: Culicidae): larval development in mixed cultures

Interactions among the larvae of Aedes aegypti (L.), Ae. albopictus (Skuse), and Ae. triseriatus (Say) were studied in trispecific and bispecific mixed populations under laboratory and field conditions. Competitive stress (as evidenced by the average time to first, 50, and 75% pupation and the total pupation periods for mixed populations of each species in comparison with their single species controls) was more pronounced in mixed cultures reared in glass jars in the laboratory than in tires under field conditions. In the laboratory, the larval development of Ae. aegypti reared together with Ae. albopictus or Ae. triseriatus, or both, larvae was accelerated significantly. Conversely, the time to pupation for Ae. albopictus and Ae. triseriatus was delayed when reared with Ae. aegypti. However, the average wing length of female Ae. albopictus and Ae. triseriatus was greater in the mixed cultures than in single species cultures. These data indicated that the effect of intraspecific competition was greater than interspecific competition. Adequate food and higher temperature appeared to promote rapid development and higher survival of the immature stages of the three Aedes species in tires placed in the field. In general, Ae. triseriatus larvae required a longer period for larval development and had greater larval mortality than either Ae. aegypti or Ae. albopictus. In mixed populations of Ae, albopictus and Ae. aegypti in the jars and food-rich tires, the periods needed to attain first, 50, and 75% cumulative pupation were not significantly different than in single species controls. We conclude that no clear-cut displacement occurred in mixed experimental populations of Ae. aegypti and Ae. albopictus.     

Ribosomal DNA structure in Aedes mosquitoes (Diptera: Culicidae) and their cell lines

The molecular organization of the ribosomal DNA repeat unit in laboratory colonies of Aedes albopictus (Skuse) and Aedes aegypti (L.) and in continuous cell lines derived from these species was examined using restriction enzyme mapping and Southern blotting techniques. In the rRNA coding regions, restriction enzyme sites were conserved in Ae. aegypti and Ae. albopictus. In particular, sites that had been mapped in rDNA cloned from the Mauritius strain of Ae. albopictus mosquitoes were confirmed in cloned DNA from cultured Ae. alboptictus cells. The intergenic spacer region of the ribosomal DNA repeat unit in Ae. albopictus was longer than that in Ae. aegypti; contained multiple recognition sites for AluI, SstI, PvuI, and XhoI; and was more heterogeneous in DNA from cultured cells than in DNA from inbred laboratory mosquitoes. The rDNA copy number was 3- to 4-fold lower in cell lines from both Ae. albopictus and Ae. aegypti than in the corresponding adult mosquitoes.     

Molecular and morphological characterization of Aedes albopictus in northwestern Greece and differentiation from Aedes cretinus and Aedes aegypti

The presence of Aedes albopictus (Skuse) was recently confirmed for the first time in northwestern Greece. This location is within the distribution range of a morphologically similar species, Aedes cretinus Edwards, and is a potentially favorable region for the reintroduction of Aedes aegypti (L.). It was thus compelling to use methods in addition to morphology-based keys to correctly identify specimens badly damaged, rubbed, or otherwise altered in their external characteristics. It was decided to use molecular techniques as a novel and reliable method for differentiating the three Stegomyia species. The nuclear internal transcribed spacer 2 (ITS2) fragments from morphologically identified Ae. albopictus and Ae. cretinus specimens were amplified, and their sequences were compared with those in GenBank for Ae. albopictus, Ae. cretinus, and Ae. aegypti. Also, mitochondrial cytochrome oxidase I (COI) fragments were amplified for Ae. albopictus and Ae. cretinus (so far not available in GenBank) and compared with Ae. aegypti fragments. ITS2 and COI sequences generated in our study were deposited in GenBank and could be useful in future studies of mosquitoes by other research workers.     

Comparative potential of Aedes triseriatus, Aedes albopictus, and Aedes aegypti (Diptera: Culicidae) to transovarially transmit La Crosse virus

Aedes triseriatus (Say) (Diptera: Culicidae), the major vector of La Crosse (LAC) virus, efficiently transmits LAC virus both horizontally and transovarially. We compared the vector competence and transovarial transmission ability of Ae. triseriatus, Aedes albopictus Skuse, and Aedes aegypti (L.) for LAC virus. Ae. triseriatus and Ae. albopictus were significantly more susceptible to oral infection with LAC virus than Ae. aegypti. The three species also differed in oral and disseminated infection rates (DIRs). Transovarial transmission (TOT) rates and filial infection rates (FIRs) were greater for Ae. triseriatus than either Ae. albopictus or Ae. aegypti. These measures were integrated into a single numerical score, the transmission amplification potential (TAP) for each species. Differences in TAP scores were due mainly to the differences in DIRs and FIRs among these mosquitoes. Although the TAP score for Ae. albopictus was lower than that of Ae. triseriatus, it was 10-fold greater than that for Ae. aegypti.     

Factors influencing ingestion of particulate materials by mosquito larvae (Diptera: Culicidae)

Relative ingestion rates of mosquito larvae, as indicated by the number of substrate-filled gut segments per unit time, were determined for Culex tarsalis Coquillett, Aedes aetypti L., and Anopheles albimanus Wiedemann. Among the three species, Ae. aegypti larvae were the most rapid feeders. F50 (median time for complete repletion of 50% of the larvae) was 61, 42, and 100 min for the three species feeding on a wheat flour suspension, respectively. Food particles with nutritive values (dried yeast, wheat flour, fishmeal, or dried blood) were ingested faster than inert particles (kaolin, talc, chalk, or charcoal). The addition of aqueous yeast extracts containing phagostimulants accelerated the ingestion of inert particles. Increasing the concentration of inert particles did not increase ingestion rates. Larval age, water temperature, and starvation, but not larval density, influenced rates of ingestion. Younger instars were more rapid feeders than older instars. First instars of the three species filled their guts with wheat flour approximately two times faster than fourth instars. Increasing water temperature from 18 to 31 degrees C accelerated wheat flour ingestion by fourth instars of Cx. tarsalis, Ae. aegypti, and An. albimanus by factors of 1.9, 1.5, and 1.7, respectively. After starvation for 12 h, fourth instars of Cx. tarsalis and Ae. aegypti increased ingestion of wheat flour about 1.6 and 1.8 times, respectively. In contrast, starvation of An. albimanus larvae for the same period resulted in decreased wheat flour ingestion by 2.2 times when compared with unstarved larvae. These results indicare that, in addition to the chemical factors associated with the food substances, the physiological and environmatal conditions of the larvae play an important role in regulating the ingestion rate of suspended particles by mosquito larvae.     

Differences in male mating response and female flight sounds in Aedes aegypti and Ae. albopictus (Diptera: Culicidae).

Aedes aegypti (L.) and Ae. albopictus (Skuse) showed similar rates and timing of insemination in the laboratory. Laboratory attempts at interspecific mating were unsuccessful. Because Ae. aegypti males are known to locate females by flight sounds, male response to sound was compared in the two species. Ae. aegypti males responded to female flight sounds with stereotypical orientation and mating behavior, whereas Ae. albopictus males seldom responded. Recorded flight sounds of females were sampled via computer digitization and compared. Ae. aegypti females produced louder sounds with more harmonics than Ae. albopictus. Males were tested for their ability to discriminate between the sounds of the two species. Ae. albopictus males did not respond to recordings of either Ae. albopictus or Ae. aegypti females. Ae. aegypti males responded preferentially to the recorded sounds of Ae. aegypti females. Thus, males of the two species use different mechanisms in locating mates. Ae. aegypti males rely more on sound than do Ae. albopictus males.     

Detritus type alters the outcome of interspecific competition between Aedes aegypti and Aedes albopictus (Diptera: Culicidae)

Many studies of interspecific competition between Aedes albopictus (Skuse) and Aedes aegypti (L.) (Diptera: Culicidae) larvae show that Ae. albopictus are superior resource competitors to Ae. aegypti. Single-species studies indicate that growth and survival of Ae. albopictus and Ae. aegypti larvae are affected by the type of detritus present in containers, which presumably affects the amount and quality of microorganisms that the mosquito larvae consume. We tested whether different detritus types alter the intensity of larval competition by raising 10 different density/species combinations of Ae. albopictus and Ae. aegypti larvae under standard laboratory conditions, with one of four detritus types (oak, pine, grass, or insect) provided as a nutrient base. Intraspecific competitive effects on survival were present with all detritus types. Ae. albopictus survivorship was unaffected by interspecific competition in all treatments. Negative interspecific effects on Ae. aegypti survivorship were present with three of four detritus types, but absent with grass. Estimated finite rate of increase (lambda') was lower with pine detritus than with any other detritus type for both species. Furthermore, Ae. aegypti lambda' was negatively affected by high interspecific density in all detritus types except grass. Thus, our experiment confirms competitive asymmetry in favor of Ae. albopictus with oak, pine, or insect detritus, but also demonstrates that certain detritus types may eliminate interspecific competition among the larvae of these species, which may allow for stable coexistence. Such variation in competitive outcome with detritus type may help to account for observed patterns of coexistence/exclusion of Ae. albopictus and Ae. aegypti in the field.     

Comparative study on nocturnal behavior of Aedes aegypti and Aedes albopictus

Nocturnal behavior of nonblood-fed females of Aedes aegypti (L.) and Aedes albopictus (Skuse) was studied using an automatic recording device equipped with a photoelectric sensor. Carbon dioxide, heating, and the contrast of the black and white colors were used as attractive cues for mosquitoes. The nocturnal host-seeking activity positively correlated with the increasing light intensity in both species. Ae. aegypti was found to be more sensitive to light than Ae. albopictus. The threshold of light intensity for the activation of the nocturnal host-seeking activity was <0.1 lx (approximately 0.01 foot candle) in Ae. aegypti and >10 lx (approximately 1 foot candle) in Ae. albopictus. Complete darkness during the daytime deactivated the host-seeking activity of both species, irrespective of their increasing flight activity controlled by their intrinsic circadian rhythms. This finding suggested that visual cues are indispensable for host-seeking behavior. The eye parameter value, the product of the ommatidial diameter, and the interommatidial angle were significantly larger in Ae. aegypti than those in Ae. albopictus, indicating that the eye of Ae. aegypti is more adapted to a darker environment.     

Larvicidal activity of sterol carrier protein-2 inhibitor in four species of mosquitoes

A previous report has shown that mosquito sterol carrier protein-2 inhibitors (SCPIs) are larvicidal to larvae of the yellowfever mosquito, Aedes aegypti (L.) (J. Lipid Res. 46: 650-657, 2005). In the current study, we tested SCPI-1 in an additional four mosquito species for larvicidal activities: Culex pipiens pipiens, Anopheles gambiae, Culex restuans, and Aedes vexans. Cholesterol accumulation in SCPI-treated Ae. aegypti fourth instars was examined. SCPI-1 is lethal to all tested mosquito species, with the LC50 value ranging from 5.2 to 15 microM when treatments started at the first to third instar. However, LC50 values increase to from 5.2 to 38.7 microM in treatments started at first and fourth instar, respectively. The results indicate that the lethal effect of SCPI-1 decreases with the growth of larvae, which suggests that SCPI-1 is more effective before the larvae reach final growth period (the last instar). SCPI-1 suppressed cholesterol uptake in Ae. aegypti fourth instars, suggesting that one of the modes of action of SCPI-1 is via reduction in cholesterol absorption.     

Susceptibility of selected strains of Aedes aegypti and Aedes albopictus (Diptera: Culicidae) to chikungunya virus.

The relative susceptibility of selected strains of Aedes aegypti (L.) and Aedes albopictus (Skuse) fed on a viremic monkey to infection with chikungunya virus was determined. Infection rates were consistently higher in 10 strains of Ae. albopictus tested than in 7 strains of Ae. aegypti tested, regardless of the geographic location from which the strains originated or the dose of virus ingested. Similarly, virus dissemination rates were higher in the Ae. albopictus strains compared with the Ae. aegypti strains. For nearly all (11 of 12) strains tested of both species, groups of mosquitoes with one or more females with a disseminated infection transmitted virus by bite to weanling mice. Based on these studies, Ae. albopictus appears to be a more competent laboratory vector of chikungunya virus than does Ae. aegypti.     

Comparative laboratory study on the reaction of Aedes aegypti and Aedes albopictus to different attractive cues in a mosquito trap

The behavioral responses of Aedes aegypti (L.) and Aedes albopictus (Skuse) adults to several attractive cues, as reactions to mosquito traps, are compared in the laboratory, and differences in the primary attractive factors for both species are discussed. Target-attacking frequency of unfed Ae. aegypti females was >30 times that of unfed Ae. albopictus females under simulated conditions. Changes in the percentage of trapped mosquitoes under several attractive conditions using commercial mosquito traps showed that Ae. aegypti mosquitoes were trapped 2-3 times faster than Ae. albopictus. For Ae. aegypti, the combination of a visual cue + CO, alone enhanced attractiveness, whereas both a visual cue + CO, as well as a visual cue + octenol enhanced Ae. albopictus. The combination of at least three factors, such as a visual cue, CO2, and a chemical cue is thought to be valuable for trapping and estimating the relative adult population sizes of Ae. aegypti and Ae. albopictus     

Reproductive and metabolic differences between Aedes aegypti and Ae. albopictus (Diptera: Culicidae).

Aedes albopictus (Skuse) females were more likely to develop eggs after they ingested small blood meals than were Aedes aegypti (L.) when both species were maintained under the same suboptimal adult nutritional regimen. The longevity of adult female Ae. albopictus under conditions of starvation was also significantly greater than that of Ae. aegypti. Analyses of total body proteins, lipids, and glycogen indicated that the increased reproductive efficiency of Ae. albopictus may be a result of its greater reserves accumulated during a longer larval period.     

Differences in energy metabolism and adult desiccation resistance among three Aedes (Stegomyia) species (Diptera: Culicidae) from South Sulawesi, Indonesia

To determine the mechanisms of adult desiccation resistance of Aedes (Stegomyia) species, we estimated the carbohydrate and lipid contents of newly emerged adult mosquitoes from South Sulawesi, Indonesia. Inter- and intraspecific differences in glycogen and free fatty acid accumulation were found in Aedes aegypti (L.), Ae. albopictus (Skuse), and Ae. paullusi (Stone & Farmer). High glycogen content was determined for both sexes in all Ae. aegypti strains and in the Ujung Pandang strain of Ae. albopictus. There were no differences among species in either trehalose or sorbitol content. A trend similar to glycogen was seen for lipid content, especially the free fatty acids of neutral lipids. These energy reserves of adult mosquitoes correlated with adult desiccation survival time at 90% RH calculated by Mogi et al. (1996). In both sexes, survival time was longer in strains with high glycogen and free fatty acid content, and low trehalose content. Ae aegypti, Ae. albopictus, and Ae. paullusi are species with high, intermediate, and low levels of glycogen and free fatty acids, respectively. However, longevity of these species under desiccation conditions is associated strongly with glycogen-trehalose conversion, and free fatty acid accumulation.     

Habitat segregation of mosquito arbovirus vectors in south Florida

Oviposition traps set in rural to urban environments in three south Florida counties were colonized predominantly by Culex quinquefasciatus Say (35.1%), Aedes albopictus (Skuse) (34.5%), Aedes aegypti (L.) (23.8%), and Culex nigripalpus Theobald (6.6%) during 1 yr of monthly sampling. Significant differences were detected among counties for abundances of Cx. quinquefasciatus and for percentage composition of that species and Ae. albopictus. Aerial images of habitats around each collection site were digitized, and coverages by each of 16 habitat variables were recorded. Abundances ofAe. aegypti were positively related to habitat variables associated with urbanization and negatively correlated to those reflecting rural characteristics. Multiple regression models of habitat selection explained similar proportions of variances in abundance of Ae. aegypti and Ae. albopictus, but signs of significant variables were opposite for these two species. No consistent trends of habitat associations were observed among counties for the two Culex spp. Co-occurrences of the four species in individual traps depended on container type (tub versus cup), and, for Aedes spp. with Culex spp., county. The results underscore the importance of scale in evaluating habitat selection and the utility of quantifiable habitat characteristics of intermediate scale to identify site characteristics favored by the arboviral vectors Ae. aegypti and Ae. albopictus.     

Variation in cold hardiness of nondiapausing eggs of nine Aedes (Stegomyia) species (Diptera: Culicidae) from eastern Asia and Pacific islands ranging from the tropics to the cool-temperate zone

Variation in cold hardiness was studied in the laboratory for the nondiapausing eggs of nine Aedes (Stegomyia) species from eastern Asia and Pacific islands, ranging from the tropics to the cool-temperate zone. Species included were Aedes aegypti (L.), Aedes albopictus (Skuse), Aedes flavopictus Yamada, Aedes galloisi Yamada, Aedes hensilli Farner, Aedes palauensis Bohart, Aedes paullusi Stone and Farner, Aedes riversi Bohart and Ingram, and Aedes scutellaris (Walker). For species with distributions covering two or three climatic zones, at least one population was included from each zone, except for Ae. aegypti (subtropical populations were unavailable). Some eggs of four species confined to the tropical zone survived short exposure to freezing temperature (> or = -5 degrees C) when humidity was high. There were inter- and intraspecies variations in cold hardiness of nondiapausing eggs. Cold hardiness in each climatic zone differed in that populations of species with more northerly distributions were more cold-hardy than species with southerly distributions. In Ae. albopictus and Ae. flavopictus that cover three climatic zones, populations from cooler regions were more cold-hardy than populations from warmer regions. A possibility of southern species without diapausing eggs to colonize temperate eastern Asia was discussed.     

Convergent habitat segregation of Aedes aegypti and Aedes albopictus (Diptera: Culicidae) in southeastern Brazil and Florida

During the rainy season of 2001, the incidence of the dengue vectors Aedes aegypti and Ae. albopictus was examined in different habitats of two cities (Rio de Janeiro and Nova Igua?u) in Rio de Janeiro State, Brazil, and in two cities (Palm Beach and Boca Raton) in Florida. Oviposition trap collections were performed in urban, suburban, and rural habitats in both areas. Our hypothesis that the abundances and frequencies of occurrence of Ae. aegypti and Ae albopictus are affected in opposite ways by increasing urbanization was only partially supported. City, habitat, and their interaction significantly affected the abundance of both species. Cities with high abundance of Ae. aegypti also had a high abundance of Ae. albopictus. The two species were most abundant in the cities of Rio de Janeiro state and the lowest in Boca Raton. Habitat had a significant but opposite effect on the abundances of Ae. aegypti and Ae. albopictus. In general, Ae. aegypti was most prevalent in highly urbanized areas and Ae. albopictus in rural, suburban, and vegetated urban areas in Rio de Janeiro state and Florida. However, abundances of the two species were similar in most suburban areas. Analyses of frequencies of occurrence showed an unexpected high level of co-occurrence of both species in the same oviposition trap. Despite the different geographical origins of Ae. albopictus in Brazil and the United States, the habitats used by this recent invader are remarkably similar in the two countries.     

Movement of dengue vectors between the human modified environment and an urban forest in Rio de Janeiro

The movement of Aedes aegypti (L.) and Aedes albopictus (Skuse) females between sylvatic and urban environments was investigated by marking, releasing, and recapturing adults and by identifying rubidium (Rb)-marked eggs of females that were released after taking a bloodmeal containing RbCI. When released in the forest, Ae. albopictus females flew as far as 1000 m and reached houses within 1 wk. When Ae. albopictus were released close to houses, most females were recaptured near the release point, and Rb-marked eggs were found 1000 m away in the forest only once, 35 d after the release. These differing patterns of movement may suggest a preference of Ae. albopictus for the human-modified environment. Ae. aegypti, however, showed low tendency to disperse into the forest. The capacity of Ae. albopictus females to disperse from a sylvatic into a human-modified environment suggests that this species may play a role in the dissemination of forest-restricted pathogens, such as yellow fever virus.