Fitness and habitat segregation of British tree-hole mosquitoes

Abstract.

• 1 Populations of three mosquitoes, Aedes geniculutus (Olivier), Anopheles plumbeus Stephens and Culex torrentium Martini, were found in three types of tree holes: rot holes penetrating the bark of the host tree and exposed, or deeper, more cryptic, bark-lined buttress holes. Aedes geniculatus occupied all three tree-hole types but reached their greatest abundance in exposed buttress holes; A.plumbeus predominated in rot holes; C.torrentium occupied buttress holes exclusively.
• 2 After a 1 month period of winter freezing, larvae of A.plumbeus survived better than larvae of A. genicularus. Freeze resistance of A.plumbeus increased from cryptic buttress to rot holes. Freeze resistance of A.geniculatus declined from exposed buttress to cryptic buttress to rot holes and was markedly higher in more protected than in more exposed macrohabitats.
• 3 Among manipulated larval cohorts of A.geniculutus and A.plumbeus, survivorship, pupation success, pupal weight, and/or biomass yield did not differ among tree-hole types or was the reverse of freeze-related survivorship. Despite seasonal differences in fitness correlates, annual fitness may be similar among tree-hole types for both species. Manipulations carried out at a single census or during a single season are therefore likely to produce misleading information about fitness variation among sub-habitats.
• 4 Both species should exploit a broad range of tree-hole types but A.plumbeus occupies more restricted types of tree holes than does A.geniculutus. The restricted habitat usage by A.plumbeus does not parallel their fitness among tree-hole types in southern Britain but does resemble that of other tree-hole anophelenes. We therefore propose that habitat selection by A.plumbeus is more likely due to their phylogenetically determined physiological tolerances and behaviour than to competition from coexisting A.geniculatus.

     

Seasonal availability of resources and habitat degradation for the western tree-hole mosquito, <Emphasis Type="Italic">Aedes sierrensis</Emphasis>

The nutrient base of aquatic tree-hole communities is derived from leaf litter, benthic detritus, and water flowing down the tree trunk (stemflow water). Previous studies in eastern North America with the mosquito, Aedes triseriatus, have identified leaf litter as a major and stemflow water as a minor source of mosquito nutrition, but did not consider the role of the benthic detritus or how the aggregate or relative contribution of these sources of mosquito nutrition changed during the year. We use the leaf litter, benthic detritus, and stemflow water from tree holes in western Oregon (USA) to determine how these substrates affect mass at metamorphosis, biomass yield, and fitness (cohort replacement rate; R ₀) of the mosquito, Aedes sierrensis, through both natural and simulated winters, the normal growing season for larvae in tree holes. We found that fresh leaf litter constitutes the major determinant of mosquito fitness by a factor of >15:1 over any other substrate taken directly from tree holes in nature. The other substrates, including the benthic detritus, individually make only a meager contribution to mosquito fitness but, when added to the leaf litter, can sustain yield and improve fitness at high, limiting larval densities. Nutritional quality of tree-hole substrates declines by >90% from early (fall) to late (spring) in the larval growing season. At both times of year, the coarse or fine detritus provide minor resources, and stemflow water provides no detectable contribution to mosquito nutrition. The resources in the litter are not transported during the year to the benthic detritus; rather, these resources are either exploited by mosquitoes when they first become available, or they deteriorate and become progressively more unavailable to them. Growth and development of A. sierrensis feeding on dried and reconstituted tree-hole contents during a 6-month simulated winter in the laboratory showed: (1) the same relative contributions of leaf litter, benthic detritus, and stemflow water to mosquito nutrition, (2) that the winter deterioration of substrate quality is a direct consequence of microbial decomposition, and (3) that pre-emptive competition from pre-existing A. sierrensis greatly increases substrate deterioration. We conclude that the progressive winter deterioration of larval resources in combination with the dry summers of western North America are the most likely environmental factors that limit species diversity in tree holes and that have selected for early recruitment (autumnal hatching) of A. sierrensis and for its univoltine life cycle from Mexico to Canada. Received: 28 December 1999 / Accepted: 10 April 2000     

Productivity of natural and artificial containers for Aedes polynesiensis and Aedes aegypti in four American Samoan villages

Six mosquito species were identified in a survey of containers associated with 347 households in four villages in American Samoa. Aedes polynesiensis Marks (Diptera: Culicidae) and Aedes aegypti (L) were the most abundant species, representing 57% and 29% of the mosquitoes identified. Culex quinquefasciatus (Say), Culex annulirostris (Skuse), Aedes oceanicus (Belkin) and Toxorhynchites amboinensis (Doleschall) were also found. Aedes aegypti and Ae. polynesiensis showed distinct differences in their use of containers, preferring large and small containers, respectively. By contrast with previous studies, Ae. polynesiensis utilized domestic and natural containers with equal frequency, whereas Ae. aegypti continued to be found predominantly in domestic containers. Only 15% of containers holding immature mosquitoes included pupae and fewer than 10 Aedes spp. pupae were found in most containers with pupae. An estimated 2289 Ae. polynesiensis and 1640 Ae. aegypti pupae were found in 2258 containers. The presence of both species in the same container did not affect the mean density of either species for larvae or pupae. Glass jars, leaf axils, tree holes and seashells produced few Aedes spp. pupae in any of the study villages. Overall, 75% of Ae. polynesiensis pupae were found in buckets, ice-cream containers and tyres, with <7% being produced in natural containers, whereas 82% of Ae. aegypti pupae were found in 44-gallon (US) drums ( approximately 166L), buckets and tyres. Source reduction efforts targeting these container types may yield significant reductions in both Ae. polynesiensis and Ae. aegypti populations in American Samoa.     

Drought and the organization of tree-hole mosquito communities

Summary In southeastern North America (North Florida, USA), the duration, frequency, and timing of drought differentially affect the survivorship of pre-adult tree-hole mosquitoes. Drought affects survivorship both by the direct action of dehydration on developing larvae and pupae and by the indirect modulation of predation. The drought-susceptible species, Toxorhynchites rutilus, Orthopodomyia signifera, and Anopheles barberi co-occur in more permanent holes that are larger, with larger, more vertical openings, lower down in larger trees, and contain darker water with higher conductivity, pH, and tannin-lignin content than the holes occupied by Aedes triseriatus that has drought-resistant eggs and rapid larval development. Ovipositing mosquitoes cue on physical and chemical attributes of tree holes independently of host tree species. These same attributes differ among drought-prone and drought-resistant holes but mosquitoes track these attributes more faithfully than the attributes predict tree-hole stability.     

Mosquitoes of Rarotonga,Cook Islands: A survey of breeding sites

Abstract

Two surveys of Rarotonga, Cook Islands (21°20^'S, 160°16^'W) were made to determine the mosquito fauna of the island, and to identify the habitats required for breeding by searching for larvae. The first survey was made during the “dry season” in May 2001, the second during the “wet season” in February 2002. The mosquito fauna comprised four species Culex (Culex) quinquefasciatus Say, Culex (Culex) annulirostris Skuse, Aedes (Stego‐myia) aegypti (Linnaeus) and Aedes (Stegomyia) polynesiensis Marks. Larvae of the Culex species were most often found in larger natural and artificial water bodies. The Aedes species bred in both natural and artificial containers of all sizes. Ae. polynesiensis was the most widespread species, using natural holes in all regions as well as artificial containers in the urban areas. Most larvae of Ae. aegypti were located in small artificial containers. The two Aedes species are the vectors of dengue fever on the island. Mosquito control during outbreaks should specifically target the artificial containers preferred by Aedes sp. for breeding habitats.     

Tree holes in a mixed broad-leaf?podocarp rain forest, New Zealand

Despite the ecological importance of tree holes as habitat for many species in New Zealand, few studies have quantified the abundance, distribution or structural characteristics of tree holes in native forests. We recorded a total of 364 tree holes in ground-to-canopy surveys on 50 trees of five endemic species in the families Fagaceae and Podocarpaceae within Orikaka Ecological Area, Buller District, New Zealand. Tree holes were not uniformly distributed throughout the forest, with more holes in the three podocarp species Prumnopitys ferruginea, P. taxifolia and Dacrycarpus dacrydioides than in Nothofagus fusca or N.?menziesii. However, N. fusca had the largest tree holes of any of the tree species sampled. Tree-hole volume and tree-hole opening both increased with tree size. Tree-hole opening was strongly positively correlated with internal volume, especially for the podocarps. It therefore potentially provides a useful surrogate measure for quantifying the abundance of large tree holes from cost-effective ground surveys. We estimated there might be as many as 771 tree holes per hectare of forest at this site, but fewer of these were thought to be large enough for obligate hole-dwelling vertebrate species. Our findings differ from those of previous researchers in native forests around New Zealand because earlier studies use ground-based surveys to sample the abundance of tree holes suitable for specific fauna, whereas we used climbed inspections where all tree holes were recorded. This could be particularly important for the conservation management of native vertebrate and invertebrate species.     

Effect of leaf litter and density on fitness and population performance of the hole mosquito Aedes triseriatus

Abstract.

• 1 We tested the effects of food, in the form of leaf litter, and density on population growth and fitness correlates of the tree hole mosquito Aedes triseriatus. Our field experiment used a 2 x 2 randomized block design, with three holes as blocks. In cages within three holes, we manipulated densities of Aedes triseriatus larvae, and presence of leaf litter. Our laboratory experiment used a 2 x 3 factorial design in which we also manipulated densities of larvae and availability of leaf litter within similar cages (inside, outside, or absent). For both experiments we determined effects on survival, days to and mass at eclosion of both sexes, and a composite index of population performance λ′, that estimates finite rate of increase.
• 2 In the field experiment, we found significant and large effects of leaf litter and of density on the performance of A. triseriatus. The effect of density did not depend on leaf litter availability. We documented significant variation among tree holes for most correlates of fitness. Often, the effects of treatments varied significantly depending on the tree hole tested.
• 3 In the laboratory experiment we found significant effects of density and litter, and that direct browsing on leaf litter is necessary for the effect of litter on performance of A. triseriatus. In the laboratory, the negative effect of increased density was dependent on leaf litter availability.
• 4 Our results demonstrate that leaf litter has important effects on population performance of tree hole mosquitoes in natural tree holes. Direct browsing on the surface of leaf litter is the most likely mechanism by which litter enhances population growth. The impact of litter on populations varies among tree holes.

     

Horizontal distribution affects the vertical distribution of native and invasive container‐inhabiting Aedes mosquitoes within an urban landscape

The vertical dimension constitutes an important niche axis along which mosquitoes may adjust their distribution. Here, we evaluated whether the vertical distribution of container‐inhabiting Aedes mosquitoes differs along a gradient of anthropogenic land‐use intensity within an urban landscape. Using a pulley system, we hung oviposition cups at three heights (ground level, 4.5, and 9 m) and in three habitats: forest, park, and a built environment. We hypothesized that mosquito abundance and diversity would be highest in the least disturbed forest habitat, decrease in the park, and be lowest at the UNC‐Greensboro campus. We also expected Aedes albopictus (Skuse) and Ae. triseriatus (Say) to mainly oviposit at ground level and Ae. hendersoni (Cockerell) at canopy height. Aedes albopictus was the most common species (68.8%) collected in all three habitat types and was the only species found in the built environment. In that habitat, Ae. albopictus exhibited a bimodal distribution with the lowest activity at the intermediate height (4.5 m). Aedes triseriatus (28.9%) did not differ in egg abundance between the forest and park habitats but did exhibit diverse vertical habitat use while avoiding the canopy in the park habitat. Aedes hendersoni (2.3%) was the most sylvatic species and oviposited only at ground level. Our results indicate that the vertical distribution of mosquitoes is affected by the type of habitat in which they occur, and that this variation could be driven via local‐scale modification of microclimatic factors.     

The first report of Aedes (Stegomyia) albopictus in Haiti

Aedes albopictus was found in six of the 10 departments of Haiti and in 14 of the 35 communes surveyed. The survey found the larvae of Ae. albopictus in 13 different types of containers. Used tires and tins were by far the most common breeding sites used by this mosquito species. At the breeding sites, Ae. albopictus was associated with other mosquito species, such as Aedes aegypti, Culex nigripalpus and Aedes mediovittatus. The highest proportion of association was with Ae. aegypti. This study represents the first report of Ae. albopictus in Haiti.     

Geographic and ecological distribution of the dengue and chikungunya virus vectors Aedes aegypti and Aedes albopictus in three major Cameroonian towns

Aedes albopictus (Diptera: Culicidae) was first reported in Central Africa in 2000, together with the indigenous mosquito species Aedes aegypti (Diptera: Culicidae). Because Ae. albopictus can also transmit arboviruses, its introduction is a public health concern. We undertook a comparative study in three Cameroonian towns (Sahelian domain: Garoua; equatorial domain: Douala and Yaoundé) in order to document infestation by the two species and their ecological preferences. High and variable levels of pre‐imaginal Ae. aegypti and Ae. albopictus infestation were detected. Only Ae. aegypti was encountered in Garoua, whereas both species were found in Douala and Yaoundé, albeit with significant differences in their relative prevalence. Peridomestic water containers were the most strongly colonized and productive larval habitats for both species. No major differences in types of larval habitat were found, but Ae. albopictus preferentially bred in containers containing plant debris or surrounded by vegetation, whereas Ae. aegypti tended to breed in containers located in environments with a high density of buildings. These findings may have important implications for vector control strategies.     

The mosquitoes of eastern Tennessee: studies on abundance,habitat preferences,and host‐seeking behaviors

A significant increase in the number of reported cases of La Crosse virus (LACV) infections in eastern Tennessee has occurred in the last ten years. The objective of this study was to determine the abundance and habitat preferences of the potential vectors of LACV in this region. Adult host‐seeking mosquitoes were collected using CO₂‐baited CDC light traps and a series of human‐landing catches in eastern Tennessee from 2004 to 2006. A total of 4,200 female mosquitoes of 23 species was collected by CO₂‐baited CDC trapping at ten sites during the study period. Aedes albopictus (Skuse) was the most abundant mosquito collected at all sites and vegetation types, with the ratios of total Ae. albopictus to Ae. triseriatus (Say) females collected being 2.1:1 in 2004, 3.8:1 in 2005, and 4.9:1 in 2006. Ten species were collected during a series of human‐landing catches made at four different sites; one probable and three confirmed case sites of LACV infections, totaling 528 female mosquitoes. Aedes albopictus was the most abundant species collected, with a 4:1 ratio of Ae. albopictus to Ae. triseriatus females. Aedes albopictus exhibited two clear peaks of “landing” activity, one in the early morning and one in the late afternoon or early evening. Simple and multiple regression analyses of the predictors of the number of mosquitoes collected showed that populations of Ae. albopictus were three times more likely to be collected overall than Ae. triseriatus. Species (Ae. albopictus), vegetation (residential), and the previous cumulative precipitation for the four weeks prior to collection were significantly (P < 0.05) associated with the number of mosquitoes collected by CO₂‐baited CDC trapping. Aedes albopictus was also more likely to be collected than Ae. triseriatus at confirmed cases of LACV infections.     

The ecology of mosquito larvae (Diptera: Culicidae) in the subalpine zone of the eastern Sierra Nevada Mountains,California, U.S.A.

Mosquito larvae were collected from the subalpine region of the eastern Sierra Nevada Mountains from 2011 to 2014. Two watersheds were sampled and sites selected were mainly vernal snow‐melt pools and wet meadows. Seven Aedes species, Culiseta incidens (Thomson), and Culex tarsalis Coquillett were collected. The most abundant and widely distributed species were Ae. hexodontus Dyar and Ae. tahoensis Dyar. Aedes tahoensis was the predominate species in woodland snow‐melt habitats. Some species were found at most elevations while others were found more often at specific elevations. The most restrictive species was Ae. ventrovittis Dyar which occurred almost exclusively between 3,219 m a.s.l. and 3,390 m a.s.l. Shannon and Simpson species diversity indices demonstrated that species diversity was greater in meadow habitats compared to woodland habitats. Mixed woodland/meadows, rock pools, and shallow grass pools were intermediate in species diversity. Abiotic factors such as snowpack and water temperature impacted species development times and when habitats dried. It was concluded that spatial and temporal patterns of habitats, along with elevation, influenced species presence and larval development. The results of the present study and previous work in the eastern Sierras will help guide future research that focuses on the potential change in the distribution and seasonality of subalpine mosquitoes and disease potential in the eastern Sierras as climatic conditions change.     

Entomological surveillance,spatial distribution,and diversity of Culicidae (Diptera) immatures in a rural area of the Atlantic Forest biome,State of São Paulo,Brazil

Because of the high adaptive capacity of mosquitoes, studies that focus on transitional environments become very important, such as those in rural areas, which are considered as bridges between wild diseases and human populations of urban areas. In this study, a survey of the existing species of mosquitoes was performed in an Atlantic Forest area of the city of Santa Bárbara d'Oeste, São Paulo state, Brazil, using traps for immatures and analyzing the frequency and distribution of these insects over the sampling months. Five mosquito species were found: Aedes albopictus (the most frequent species), Aedes aegypti, Aedes fluviatilis, Culex quinquefasciatus, and Toxorhynchites theobaldi. The 4,524 eggs collected in ovitraps showed the presence of the tribe Aedini. Aedes aegypti and Ae. albopictus were identified after larval hatching in the laboratory, with different spatial distributions: the first of which coincides with the area of greatest diversity calculated using the Simpson index, while the second does not. The association of ecological analysis of spatial diversity with simple methods of data collection enables the identification of possible epidemiological risk situations and is a strategy that may be implemented to monitor ecological processes resulting from the interaction among different species of mosquitoes.     

Seasonal and diel patterns of biting midges (Ceratopogonidae) and mosquitoes (Culicidae) on the Parris Island Marine Corps Recruit Depot

The Marine Corps Recruit Depot on Parris Island, SC, is surrounded by tidal salt marshes, which are breeding habitats for many pestiferous biting flies. Knowledge of biting fly behavior patterns is needed to develop effective pest management strategies in urban areas adjacent to salt marshes. We measured biting midge (Ceratopogonidae) and mosquito (Culicidae) seasonal abundance and diel activity patterns on Parris Island using CO₂‐baited suction traps from November 2001 – November 2004. Of the three biting midge species collected, Culicoides furens was most abundant (86.2% of total) and was present in high numbers from late March to November. Culicoides hollensis (12.0% of total) was present during spring and fall but absent in summer and winter; and Culicoides melleus (1.7% of total) was present in spring through fall but absent in winter. Abundance of C. furens had a positive linear correlation with air temperature and rainfall. There were nonlinear correlations between air temperature and C. hollensis and C. melleus numbers, which were most abundant at moderate temperatures. Of 18 mosquito species collected, the most abundant were Aedes taeniorhynchus (42.7% of total), Aedes sollicitans (26.3% of total), Culex salinarius (15.6% of total), Culex quinquefasciatus (7.3% of total), and Aedes vexans (5.7% of total); other species comprised <5% of collections. Aedes taeniorhynchus numbers were positively correlated with temperature and rainfall, and Cx. salinarius was correlated with soil moisture. Activity of most biting midges and mosquitoes were highest the first two hours following sunset. Species of biting flies were present in all months, suggesting that year‐round control measures are necessary to reduce exposure to potential disease vectors and nuisance biting.     

Predator-mediated,non-equilibrium coexistence of tree-hole mosquitoes in southeastern North America

Summary Mosquito populations in tree holes in northern Florida (30.6° N lat.), USA are held below their carrying capacities by a self-limiting, cannabalistic predator. Within tree holes, extinctions and reinvasions are common; in the system as a whole, extinctions and immigrations occur without regard to community composition, tree-hole size or stability, or average number of species present. Little, if any, density-dependent development takes place. There is no evidence that the community ever reaches equilibrium, that competition is taking place, or that competition has been an important factor structuring this mosquito community. Rather, examination of related species in the same genera suggests that the principal determinants of their coexistence relate to the adaptations already possessed by each species at the time of their first encounter. Thus, unless experimentally demonstrated or reasonably inferred from circumstantial evidence, competition and coevolved niche shifts cannot be invoked to explain the coexistence of a diversity of species within a habitat type, no matter how circumscribed or discrete that habitat.     

First record and establishment of the mosquito Aedes albopictus in Spain

The invasive mosquito Aedes (Stegomyia) albopictus (Skuse) (Diptera: Culicidae) was detected for the first time in Spain, in Sant Cugat del Vallès, a city in the north-east of the country (41 degrees 28' N, 2 degrees 4' E, altitude 120 m), during August 2004. A male and one larva were collected in the backyard of a house and in a tree hole, respectively. Dense populations of adults and larvae were found in subsequent surveys, confirming the establishment of the species in the area. This is the first report of the establishment of this species in the Iberian Peninsula.     

Forest canopy,water level,and biopesticide interact to determine oviposition habitat selection in Aedes albopictus

Understanding how interacting abiotic and biotic factors influence colonization rates into different habitat types is critical for both conserving and controlling species. For example, the rapid global spread of Asian tiger mosquitoes, Aedes albopictus, has reduced native species abundances and produced disease outbreaks. Fortunately, bacterial endospores of two Bacillus species (biospesticide) are highly lethal to Ae. albopictus larvae and have been commercially developed to reduce populations. Oviposition habitat selection is the first defense Ae. albopictus females possess against any control substance added to breeding sites, and considerable variation exists in their response to biopesticides. In a field experiment, I crossed the presence/absence of biopesticides, with two canopy (open, closed) and water (high, low) levels at 64 breeding sites, to examine if these interacted to influence oviposition site choice. Avoidance of biopesticide was most pronounced in closed canopy sites and those with low water levels, as all main effects and two‐way interactions influenced oviposition. Oviposition habitat selection represents a possible mechanism of resistance to biopesticides and other methods used to kill mosquito larvae. Future experiments examining how larval density and mortality modify these results should allow for more effective control of this highly invasive species.     

Worldwide patterns of genetic differentiation imply multiple 'domestications' of Aedes aegypti, a major vector of human diseases

Understanding the processes by which species colonize and adapt to human habitats is particularly important in the case of disease-vectoring arthropods. The mosquito species Aedes aegypti, a major vector of dengue and yellow fever viruses, probably originated as a wild, zoophilic species in sub-Saharan Africa, where some populations still breed in tree holes in forested habitats. Many populations of the species, however, have evolved to thrive in human habitats and to bite humans. This includes some populations within Africa as well as almost all those outside Africa. It is not clear whether all domestic populations are genetically related and represent a single 'domestication' event, or whether association with human habitats has developed multiple times independently within the species. To test the hypotheses above, we screened 24 worldwide population samples of Ae. aegypti at 12 polymorphic microsatellite loci. We identified two distinct genetic clusters: one included all domestic populations outside of Africa and the other included both domestic and forest populations within Africa. This suggests that human association in Africa occurred independently from that in domestic populations across the rest of the world. Additionally, measures of genetic diversity support Ae. aegypti in Africa as the ancestral form of the species. Individuals from domestic populations outside Africa can reliably be assigned back to their population of origin, which will help determine the origins of new introductions of Ae. aegypti.