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.     

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.     

Interspecific larval competition between Aedes albopictus and Aedes japonicus (Diptera: Culicidae) in northern Virginia

Aedes albopictus (Skuse) and Aedes japonicus (Theobald) are two of the most recent and widespread invasive mosquito species to have become established in the United States. The two species co-occur in water-filled artificial containers, where crowding and limiting resources are likely to promote inter- or intraspecific larval competition. The performance of northern Virginia populations of Ae. japonicus and Ae. albopictus competing as larvae under field conditions was evaluated. Per capita rates of population increase for each species were estimated, and the effects of species composition and larval density were determined. In water-containing cups provided with oak leaves, Ae. albopictus larvae exhibited a competitive advantage over Ae. japonicus as a consequence of higher survivorship, shorter developmental time, and a significantly higher estimated population growth rate under conditions of interspecific competition. Intraspecific competition constrained population performance of Ae. albopictus significantly more than competition with Ae. japonicus. In the context of the Lotka-Volterra model of competition, these findings suggest competitive exclusion of Ae. japonicus in those habitats where this species co-occurs with Ae. albopictus.     

Geographic distribution and breeding site preference of Aedes albopictus and Aedes aegypti (Diptera: culicidae) in Cameroon, Central Africa

Presence in Cameroon of the recently introduced Aedes (Stegomyia) albopictus (Skuse) in association with the indigenous Aedes aegypti (L.) raises public heath concerns because it might alter the risk of arbovirus transmission. The breeding site and distribution of the two Stegomyia species are updated and reported following entomological surveys carried out in 22 localities throughout Cameroon, with a total of 1,353 containers with water visited. Ae. aegypti was found in every location sampled, showing higher infestation rates in northern Cameroon. Breeding populations of Ae. albopictus were observed in all 19 southern localities, up to the Adamaoua mountains, but the species was not recorded further north. In the area where both species are present, they were often sampled in the same larval developmental sites, suggesting convergent habitat segregation. The most frequently encountered artificial and natural breeding sites were used tires, discarded tins and plastic containers, abandoned car parts, brick holes, dead leaves on the ground, tree holes, and rock pools. Further monitoring of the demographic as well as geographic expansion of Ae. albopictus in this Afrotropical environment and its relationships with indigenous Ae. aegypti should provide insight into the biology of this highly invasive species and help to implement arboviruses surveillance programs in the area.     

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.     

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.     

Larval mosquito habitat utilization and community dynamics of Aedes albopictus and Aedes japonicus (Diptera: Culicidae)

Aedes albopictus (Skuse) and Ae. japonicus (Theobald) are important container-inhabiting mosquitoes that transmit disease agents, outcompete native species, and continue to expand their range in the United States. Both species deposit eggs in natural and artificial containers and thrive in peridomestic environments. The goal of our study was to examine the types and characteristics of containers that are most productive for these species in the northeastern United States. In total, 306 containers were sampled in urban, suburban, and rural areas of New Jersey. Multiple biotic and abiotic factors were recorded in an attempt to identify variables associated with the productivity of each species. Based on pupal abundance and density of container types, results showed that tires, trash cans, and planter dishes were the most important containers for Ae. albopictus, while planter dishes were the most important containers for Ae. japonicus. Container color (black and gray), material (rubber), and type (tires) were correlated with species presence for Ae. albopictus and Ae. japonicus. These factors may play a role in the selection of oviposition sites by female mosquitoes or in the survival of their progeny. Differences in species composition and abundance were detected between areas classified as urban, suburban, and rural. In urban and suburban areas, Ae. albopictus was more abundant in container habitats than Ae. japonicus; however, Ae. japonicus was more abundant in rural areas, and when water temperatures were below 14 degrees C. Our results suggest many variables can influence the presence of Ae. albopictus and Ae. japonicus in container habitats in northeastern United States.     

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     

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.     

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.     

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.     

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.     

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.     

Behavioral differences of invasive container-dwelling mosquitoes to a native predator

Aquatic prey show behavioral modifications in the presence of predation-risk cues that alleviate their risk from predation. Aedes albopictus (Skuse), Aedes japonicus (Theobald), and Culex pipiens L. are invasive mosquitoes in North America, and their larvae are prey for the native mosquito predator, Toxorhynchites rutilus (Coquillett). Ae. albopictus and Ae. japonicus are recent invaders, whereas Cx. pipiens has been in the United States for >100 yr. In the presence of predation-risk cues from Tx. rutilus larvae, Cx. pipiens larvae increased the time spent resting at the surface (least risky behavior) more than the other prey species. Ae. japonicus larvae increased resting at the surface of the containers more than Ae. albopictus larvae in the presence of predation-risk cues. Cx. pipiens larvae spent more time motionless at the surface even in the absence of predation-risk cues when compared with the other species, indicating that Cx. pipiens larvae are the least vulnerable prey. As compared with the other prey species, Ae. albopictus larvae exhibited more high-risk behaviors both in the presence and absence of predation-risk cues, indicating that they are the most vulnerable prey. Ae. albopictus is the superior competitor; however, predation by Tx. rutilus larvae may prevent competitive exclusion by Ae. albopictus and promote coexistence among the three prey species.     

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.     

Asymmetrical competition and patterns of abundance of Aedes albopictus and Culex pipiens (Diptera: Culicidae)

We tested for competitive advantage among larvae of Aedes albopictus (Skuse) and Culex pipiens L. in a laboratory experiment and determined the frequency and spatial and temporal patterns of co-occurrence in the field in East St. Louis, IL. In a laboratory competition experiment at multiple combined densities of Ae. albopictus and Cx. pipiens larvae, Ae. albopictus survivorship and developmental times were significantly affected by conspecific densities but not by Cx. pipiens densities. In contrast, Cx. pipiens survivorship and developmental times were significantly affected by both conspecific and Ae. albopictus densities. Per capita rate of increase (r') for Ae. albopictus cohorts declined significantly due to density of conspecifics, but not density of Cx. pipiens. Interspecific competition between Ae. albopictus and Cx. pipiens under these laboratory conditions was strong and asymmetrical, with the effect of Ae. albopictus on Cx. pipiens much stronger than the reverse. In monthly samples from tire sites in East St. Louis, Ae. albopictus was highly seasonal, occurring in relatively low abundance from early May to July and increasing in abundance in August and September. Co-occurrence corresponded to the seasonality of Ae. albopictus, with Cx. pipiens encountering Ae. albopictus in more tires and at higher numbers within a tire, in August and September. Abundance of both species was high in residential areas and was unrelated to overstory cover, total nitrogen, and total phosphorus. Abundance of Ae. albopictus, but not of Cx. pipiens, was positively associated with conductivity. We expect Cx. pipiens to suffer from the effects of interspecific competition in tires in which it encounters Ae. albopictus. Interspecific competition between these species may be of both ecological and medical importance.     

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.     

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.     

Thermal sensitivity of Aedes aegypti from Australia: empirical data and prediction of effects on distribution

An understanding of physiological sensitivity to temperature and its variability is important for predicting habitat suitability for disease vectors under different climatic regimes. In this study, we characterized the thermal sensitivity of larval developmental rates and survival in several Australian mainland populations of the dengue virus vector Aedes aegypti. Males developed more rapidly than females, but there were no differences among populations for development time or survival despite previously demonstrated genetic differentiation for neutral markers. Optimal development and survival temperatures were 37 degrees C and 25 degrees C, respectively. The values for maximal development and survival were similar to standard functions used in the container inhabiting simulation (CIMSIM) model for predicting population dynamics ofAe. aegypti populations, but CIMSIM assumed a lower optimal temperature. Heat stress experiments indicated that larvae could withstand water temperatures up to 44 degrees C regardless of the rate at which temperature was increased. Results from development time measured under constant temperatures could predict development time under fluctuating conditions, whereas CIMSIM predicted faster rates of development. This difference acts to reduce the predicted potential number of generations of Ae. aegypti per year in Australia, although it does not influence its predicted distribution, which depends critically on the nature of the aquatic breeding sites.     

A polymerase chain reaction-based diagnostic to identify larvae and eggs of container mosquito species from the Australian region

Dengue outbreaks occur regularly in parts of northern Queensland, Australia, and there is concern that these outbreaks may spread with the introduction and range expansion of the two main vectors Aedes aegypti (L.) and Aedes albopictus (Skuse). Problems encountered in separating larvae of endemic and exotic container mosquito species resulted in the development of a polymerase chain reaction diagnostic procedure that uses a restriction enzyme to cut the internal transcribed spacer region 1 of the ribosomal DNA to separate Ae. aegypti and Ae. albopictus from a number of common local container mosquito species which can be used at any stage of the life cycle, including eggs up to 8 wk of age. Identification was possible using desiccated or alcohol-preserved specimens with a response time of < 24 h after receipt of the specimens.