Seasonal dynamics and habitat specificity of mosquitoes in an English wetland: implications for UK wetland management and restoration

We engaged in field studies of native mosquitoes in a Cambridgeshire Fen, investigating a) the habitat specificity and seasonal dynamics of our native fauna in an intensively managed wetland, b) the impact of water‐level and ditch management, and c) their colonization of an arable reversion to flooded grassland wetland expansion project. Studies from April to October, 2010 collected 14,000 adult mosquitoes (15 species) over 292 trap‐nights and ～4,000 pre‐imaginal mosquitoes (11 species). Open floodwater species (Aedes caspius and Aedes cinereus, 43.3%) and wet woodland species (Aedes cantans/annulipes and Aedes rusticus, 32.4%) dominated, highlighting the major impact of seasonal water‐level management on mosquito populations in an intensively managed wetland. In permanent habitats, managing marginal ditch vegetation and ditch drying significantly affect densities of pre‐imaginal anophelines and culicines, respectively. This study presents the first UK field evidence of the implications of wetland expansion through arable reversion on mosquito colonization. Understanding the heterogeneity of mosquito diversity, phenology, and abundance in intensively managed UK wetlands will be crucial to mitigating nuisance and vector species through habitat management and biocidal control.     

Bacterial microbiota assemblage in Aedes albopictus mosquitoes and its impacts on larval development

Interactions between bacterial microbiota and mosquitoes play an important role in mosquitoes’ capacity to transmit pathogens. However, microbiota assemblages within mosquitoes and the impact of microbiota in environments on mosquito development and survival remain unclear. This study examined microbiota assemblages and the effects of aquatic environment microbiota on the larval development of the Aedes albopictus mosquito, an important dengue virus vector. Life table studies have found that reducing bacterial load in natural aquatic habitats through water filtering and treatment with antibiotics significantly reduced the larva‐to‐adult emergence rate. This finding was consistent in two types of larval habitats examined—discarded tires and flowerpots, suggesting that bacteria play a crucial role in larval development. Pyrosequencing of the bacterial 16S rRNA gene was used to determine the diversity of bacterial communities in larval habitats and the resulting numbers of mosquitoes under both laboratory and field conditions. The microbiota profiling identified common shared bacteria among samples from different years; further studies are needed to determine whether these bacteria represent a core microbiota. The highest microbiota diversity was found in aquatic habitats, followed by mosquito larvae, and the lowest in adult mosquitoes. Mosquito larvae ingested their bacterial microbiota and nutrients from aquatic habitats of high microbiota diversity. Taken together, the results support the observation that Ae. albopictus larvae are able to utilize diverse bacteria from aquatic habitats and that live bacteria from aquatic habitats play an important role in larval mosquito development and survival. These findings provide new insights into bacteria's role in mosquito larval ecology.     

Drug resistance to sulphadoxine-pyrimethamine in Plasmodium falciparum malaria in Mlimba, Tanzania

Background

In the past two decades the east African highlands have experienced several major malaria epidemics. Currently there is a renewed interest in exploring the possibility of anopheline larval control through environmental management or larvicide as an additional means of reducing malaria transmission in Africa. This study examined the landscape determinants of anopheline mosquito larval habitats and usefulness of remote sensing in identifying these habitats in western Kenya highlands.

Methods

Panchromatic aerial photos, Ikonos and Landsat Thematic Mapper 7 satellite images were acquired for a study area in Kakamega, western Kenya. Supervised classification of land-use and land-cover and visual identification of aquatic habitats were conducted. Ground survey of all aquatic habitats was conducted in the dry and rainy seasons in 2003. All habitats positive for anopheline larvae were identified. The retrieved data from the remote sensors were compared to the ground results on aquatic habitats and land-use. The probability of finding aquatic habitats and habitats with Anopheles larvae were modelled based on the digital elevation model and land-use types.

Results

The misclassification rate of land-cover types was 10.8% based on Ikonos imagery, 22.6% for panchromatic aerial photos and 39.2% for Landsat TM 7 imagery. The Ikonos image identified 40.6% of aquatic habitats, aerial photos identified 10.6%, and Landsate TM 7 image identified 0%. Computer models based on topographic features and land-cover information obtained from the Ikonos image yielded a misclassification rate of 20.3–22.7% for aquatic habitats, and 18.1–25.1% for anopheline-positive larval habitats.

Conclusion

One-metre spatial resolution Ikonos images combined with computer modelling based on topographic land-cover features are useful tools for identification of anopheline larval habitats, and they can be used to assist to malaria vector control in western Kenya highlands.     

Mosquitoes and other aquatic insects in fallow field biotopes and rice paddy fields

Fallow field biotopes that develop from abandoned rice fields are man‐made wetlands that provide new habitats for various aquatic animals. Although consideration of such biotopes generally focuses on their positive aspects, this study evaluated the negative aspects of establishing fallow field biotopes with regard to mosquito breeding sites. To determine whether fallow field biotopes become breeding habitats for vector mosquitoes, we evaluated mosquito fauna in fallow field biotopes and adjacent rice fields. We found larvae of Anopheles lesteri, Anopheles sinensis and Culex tritaeniorhynchus (all: Diptera: Culicidae) in the biotopes. Although abundances of mosquito larvae in the biotopes and rice fields were statistically similar, mosquito abundances in rice fields increased dramatically in August when the water level reduced after the rainy season. The abundance and variety of the mosquitoes' natural predators were greater in biotopes than in rice fields because the former are a permanent and stable aquatic environment. A generalized linear mixed model showed a negative effect of predator diversity on mosquito larvae abundance in both habitats. Although fallow field biotopes become breeding habitats for vector mosquitoes, establishing biotopes from fallow fields in order to protect various aquatic animals, including mosquito insect predators, may help to control mosquito breeding.     

Efficacy of aquatain, a monomolecular film, for the control of malaria vectors in rice paddies

Background

Rice paddies harbour a large variety of organisms including larvae of malaria mosquitoes. These paddies are challenging for mosquito control because their large size, slurry and vegetation make it difficult to effectively apply a control agent. Aquatain, a monomolecular surface film, can be considered a suitable mosquito control agent for such breeding habitats due to its physical properties. The properties allow Aquatain to self-spread over a water surface and affect multiple stages of the mosquito life cycle.

Methodology/Principal Findings

A trial based on a pre-test/post-test control group design evaluated the potential of Aquatain as a mosquito control agent at Ahero rice irrigation scheme in Kenya. After Aquatain application at a dose of 2 ml/m² on rice paddies, early stage anopheline larvae were reduced by 36%, and late stage anopheline larvae by 16%. However, even at a lower dose of 1 ml/m² there was a 93.2% reduction in emergence of anopheline adults and 69.5% reduction in emergence of culicine adults. No pupation was observed in treated buckets that were part of a field bio-assay carried out parallel to the trial. Aquatain application saved nearly 1.7 L of water in six days from a water surface of 0.2 m² under field conditions. Aquatain had no negative effect on rice plants as well as on a variety of non-target organisms, except backswimmers.

Conclusions/Significance

We demonstrated that Aquatain is an effective agent for the control of anopheline and culicine mosquitoes in irrigated rice paddies. The agent reduced densities of aquatic larval stages and, more importantly, strongly impacted the emergence of adult mosquitoes. Aquatain also reduced water loss due to evaporation. No negative impacts were found on either abundance of non-target organisms, or growth and development of rice plants. Aquatain, therefore, appears a suitable mosquito control tool for use in rice agro-ecosystems.     

Physical,Chemical and Biological Characteristics in Habitats of High and Low Presence of Anopheline Larvae in Western Kenya Highlands

Background

Characteristics of aquatic habitats determine whether mosquitoes will oviposit, hatch, develop, pupate and successfully emerge into adults or not, thus influencing which mosquito species will occupy a habitat. This study determined whether physiochemical and biological characteristics differ between habitats with high and low presence of anopheline larvae.

Methods

Physical, chemical and biological characteristics were evaluated in selected habitats twice per month within three highland valleys in western Kenya. Aquatic macro-organisms were sampled using a sweep-net. Colorimetric methods were used to determine levels of iron, phosphate, nitrate, ammonium and nitrite in water samples. Generalized Estimating Equations (GEE) was used to compare parameters between the two categories of anopheline presence.

Results

Habitats with high anopheline presence had greater abundance of mosquito aquatic stages and tadpoles and two times more levels of nitrate in water, whereas habitats with low anopheline presence had wider biofilm cover and higher levels of iron in water.

Conclusion

Habitats of high and low presence of anopheline larvae, which differed in a number of physical, chemical and biological characteristics, were identified in valleys within western Kenya highlands. Differences in habitat characteristics are critical in determining the number of anopheline larvae that will fully develop and emerge into adults.     

Aquatic microfauna alter larval food resources and affect development and biomass of West Nile and Saint Louis encephalitis vector Culex nigripalpus (Diptera: Culicidae)

Ciliate protists and rotifers are ubiquitous in aquatic habitats and can comprise a significant portion of the microbial food resources available to larval mosquitoes, often showing substantial declines in abundance in the presence of mosquito larvae. This top‐down regulation of protists is reported to be strong for mosquitoes inhabiting small aquatic containers such as pitcher plants or tree holes, but the nature of these interactions with larval mosquitoes developing in other aquatic habitats is poorly understood. We examined the effects of these two microbial groups on lower trophic level microbial food resources, such as bacteria, small flagellates, and organic particles, in the water column, and on Culex larval development and adult production. In three independent laboratory experiments using two microeukaryote species (one ciliate protist and one rotifer) acquired from field larval mosquito habitats and cultured in the laboratory, we determined the effects of Culex nigripalpus larval grazing on water column microbial dynamics, while simultaneously monitoring larval growth and development. The results revealed previously unknown interactions that were different from the top‐down regulation of microbial groups by mosquito larvae in other systems. Both ciliates and rotifers, singly or in combination, altered other microbial populations and inhibited mosquito growth. It is likely that these microeukaryotes, instead of serving as food resources, competed with early instar mosquito larvae for microbes such as small flagellates and bacteria in a density‐dependent manner. These findings help our understanding of the basic larval biology of Culex mosquitoes, variation in mosquito production among various larval habitats, and may have implications for existing vector control strategies and for developing novel microbial‐based control methods.     

Diversity of anopheline mosquitoes (Diptera: Culicidae) and classification based on the characteristics of the habitats where they were collected in Puerto Iguazú, Misiones,Argentina

In order to extend the knowledge of anopheline diversity and their habitats in three environments with different degrees of anthropic intervention in Puerto Iguazú, Misiones, anopheline larvae were collected and classified on the basis of similarities of their habitats. Spatio‐temporal abundance was determined and larval diversity and complementarity index were calculated. Rank‐abundance curves were performed to compare the composition, abundance, and species evenness among environments. A total of 783 larvae, belonging to six species: Anopheles argyritarsis, An. fluminensis, An. mediopunctatus, An. punctimacula, An. strodei s.l., and An. triannulatus s.l., were collected. A cluster analysis and a principal component analysis detected two groups; exposure to sunlight and type of habitat were the characteristics that explained the grouping of species. Higher abundances of anopheline larvae were observed during autumn and spring. The greatest richness was recorded in wild and peri‐urban environments and the effective number of species was greater in the wild. Anopheles punctimacula and An. triannulatus s.l. are secondary vectors of malaria in other South American countries and both species were found in the three environments, so that deforestation poses a potential risk for malaria transmission as it contributes to the proliferation of larval habitats for these mosquitoes.     

Mosquito fauna inhabiting water bodies in the urban environment of Córdoba city,Argentina, following a St. Louis encephalitis outbreak

An understanding of urban aquatic environments as mosquito larval habitats is necessary to prioritize sites for surveillance and control of arbovirus vectors in urban areas. Natural and artificial water bodies at ground level that may be larval mosquito habitats in Córdoba city, Argentina were surveyed. Data on the characteristics of aquatic sites and the presence and abundance of mosquito larvae and pupae were collected in the summer of 2006, coinciding with the first report of human WNV and following an outbreak of St. Louis encephalitis in 2005. Eight species in the genera Aedes, Culex, and Mansonia were identified. At 64.2% (34 of 53) of the sites, only one species was collected, while 3.8% (2 of 53) had three associated species, the highest richness found per site. Culex quinquefasciatus represented over 99% (out of 32,729) of the specimens. It was also the most widely distributed and detected under diverse habitat conditions. Although puddles and semi‐permanent pools harbored a greater number of species, drainages and channels may be more relevant as risk factors from an epidemiological point of view because they showed the highest larval densities, mainly of Cx. quinquefasciatus (vector of SLE and WNV). Also, higher densities of this species were associated with stormwater runoff and sewage water, thus water management systems should be targeted and closely monitored for mosquito control purposes.     

Effects of flash flooding on mosquito and community dynamics in experimental pools

Pulsed disturbances of larval mosquito sites are likely to have a direct negative effect on mosquitoes but may also have indirect effects due to the alteration of community structure. These altered communities may become attractive to gravid mosquitoes searching for oviposition sites when the disturbances decrease the abundance of mosquito antagonists such as competitors, which often results in an increase in mosquito food resources. However, flash flood disturbances in intermittent riverbeds can also remove mosquito food resources such as algae, so that the net effect of flash floods could be either to increase or decrease mosquito abundance. We conducted an outdoor mesocosm experiment to assess the effects of flash floods on mosquito oviposition habitat selection and larval abundance during the post‐disturbance period of community recovery. Mesocosms were artificially flooded. Mosquito oviposition, immature abundance, invertebrate species diversity, chlorophyll a, and abiotic parameters were monitored. Our results showed that the flash flood negatively affected phytoplankton and zooplankton, leading to a decrease of mosquito oviposition in flooded mesocosms compared to non‐flooded mesocosms. More broadly, this study indicates how disturbances influence mosquito oviposition habitat selection due to the loss of food resources in ephemeral pools, and it highlights the importance of considering the effects of disturbances in management, habitat restoration, and biodiversity conservation in temporary aquatic habitats.     

How do local differences in saltmarsh ecology influence disease vector mosquito populations?

Saltmarsh breeding mosquitoes are an important source of vectors for arboviral transmission. In southern Australia, the most prominent vector borne disease, Ross River virus (Togaviridae: Alphavirus) (RRV), is transmitted by the saltmarsh mosquito (Diptera: Culicidae) Aedes camptorhynchus (Thomson). However, the factors driving the abundance of this mosquito within and among saltmarshes are poorly understood. To predict the abundance of this mosquito within saltmarshes, the environmental conditions and aquatic invertebrate ecology of three temperate saltmarshes habitats were monitored over two seasons. Up to 44% of first-instar mosquito numbers and 21% of pupal numbers were accounted for by environmental variables. Samphire vegetation cover was a common predictor of first-instar numbers across sites although, between saltmarshes, aquatic factors such as high salinity, temperatures less than 22 °C and water body volume were important predictors. The identified predictors of pupal numbers were more variable and included high tides, waterbody volume and alkalinity. The composition of invertebrate functional feeding groups differed between saltmarshes and showed that an increased diversity led to fewer mosquitoes. It was evident that apparently similar saltmarshes can vary markedly in invertebrate assemblages, water availability and conditions through tidal inundations, rainfall or waterbody permanency. The present study advances insight into predictors of vector mosquito numbers that drive the risk of RRV outbreaks.     

The roles of history: age and prior exploitation in aquatic container habitats have immediate and carry‐over effects on mosquito life history

1. Per‐capita resource availability in aquatic habitats is influenced directly by consumer density via resource competition and indirectly via delayed resource competition when temporally non‐overlapping cohorts of larvae exploit the same resources. In detritus‐based systems, resources are likely to be influenced by the age of the aquatic habitat, as detritus changes in quality over time and may be replenished by new inputs. 2. For aquatic insects that exploit detritus‐based habitats, feeding conditions experienced during immature stages can influence fitness directly via effects on development and survivorship, but also indirectly by influencing adult traits such as fecundity and longevity. 3. Larval habitat age and prior resource exploitation were manipulated in a field experiment using the container mosquito Aedes triseriatus. 4. It was found that A. triseriatus from older habitats had greater larval survival, faster development and greater adult longevity. Exploitation of larval habitats by a prior cohort of larvae had a significant negative effect on subsequent cohorts of larvae by delaying development. 5. It is suggested that extended conditioning of detritus probably resulted in conversion of recalcitrant resources to more available forms which improved the quality of the habitat. 6. In a parallel study, evidence was found of carry‐over effects of habitat age and prior exploitation on adult longevity for A. triseriatus and Aedes japonicus collected from unmanipulated aquatic habitats. 7. These results indicate the importance of detritus dynamics and the discontinuous nature of resource competition in these mosquito‐dominated aquatic systems.     

Inhibition of Malaria Infection in Transgenic Anopheline Mosquitoes Lacking Salivary Gland Cells

Malaria is an important global public health challenge, and is transmitted by anopheline mosquitoes during blood feeding. Mosquito vector control is one of the most effective methods to control malaria, and population replacement with genetically engineered mosquitoes to block its transmission is expected to become a new vector control strategy. The salivary glands are an effective target tissue for the expression of molecules that kill or inactivate malaria parasites. Moreover, salivary gland cells express a large number of molecules that facilitate blood feeding and parasite transmission to hosts. In the present study, we adapted a functional deficiency system in specific tissues by inducing cell death using the mouse Bcl-2-associated X protein (Bax) to the Asian malaria vector mosquito, Anopheles stephensi. We applied this technique to salivary gland cells, and produced a transgenic strain containing extremely low amounts of saliva. Although probing times for feeding on mice were longer in transgenic mosquitoes than in wild-type mosquitoes, transgenic mosquitoes still successfully ingested blood. Transgenic mosquitoes also exhibited a significant reduction in oocyst formation in the midgut in a rodent malaria model. These results indicate that mosquito saliva plays an important role in malaria infection in the midgut of anopheline mosquitoes. The dysfunction in the salivary glands enabled the inhibition of malaria transmission from hosts to mosquito midguts. Therefore, salivary components have potential in the development of new drugs or genetically engineered mosquitoes for malaria control.     

Hemolytic C-type lectin CEL-III from sea cucumber expressed in transgenic mosquitoes impairs malaria parasite development

The midgut environment of anopheline mosquitoes plays an important role in the development of the malaria parasite. Using genetic manipulation of anopheline mosquitoes to change the environment in the mosquito midgut may inhibit development of the malaria parasite, thus blocking malaria transmission. Here we generate transgenic Anopheles stephensi mosquitoes that express the C-type lectin CEL-III from the sea cucumber, Cucumaria echinata, in a midgut-specific manner. CEL-III has strong and rapid hemolytic activity toward human and rat erythrocytes in the presence of serum. Importantly, CEL-III binds to ookinetes, leading to strong inhibition of ookinete formation in vitro with an IC(50) of 15 nM. Thus, CEL-III exhibits not only hemolytic activity but also cytotoxicity toward ookinetes. In these transgenic mosquitoes, sporogonic development of Plasmodium berghei is severely impaired. Moderate, but significant inhibition was found against Plasmodium falciparum. To our knowledge, this is the first demonstration of stably engineered anophelines that affect the Plasmodium transmission dynamics of human malaria. Although our laboratory-based research does not have immediate applications to block natural malaria transmission, these findings have significant implications for the generation of refractory mosquitoes to all species of human Plasmodium and elucidation of mosquito-parasite interactions.     

Effect of habitat complexity on the predation of Buenoa fuscipennis (Heteroptera: Notonectidae) on mosquito immature stages and alternative prey

Notonectids are well‐known predators in aquatic habitats, where mosquito larvae, chironomids, and cladocerans constitute their main diet. Our purpose was to assess the effect of structural complexity on the predatory ability of Buenoa fuscipennis, a common predator in aquatic habitats of Buenos Aires city (Argentina). Buenoa fuscipennis showed type 2 functional responses in both the presence and absence of prey refuge and no differences in attack rate or handling time between refuge treatments. Regarding mosquito size classes, B. fuscipennis exhibited a significantly higher preference for 2^nd instar larvae and no predation on pupae. In the presence of mosquito larvae and alternative prey, B. fuscipennis preferred mosquitoes over chironomid larvae and adult cladocerans over mosquito larvae. No switching behavior was detected in our experiments. Habitat structure only slightly affected the predator´s consumption rates on mosquito larvae. Overall, preference for prey did not vary with the presence of refuge, except for the preference for mosquitoes over chironomid larvae, which was significantly decreased in the presence of refuge as a consequence of reduced predation on mosquito larvae. The results suggest that B. fuscipennis could efficiently control mosquitoes in structurally simple habitats where chironomids are the most abundant alternative prey but not in temporary pools where cladocerans are abundant.     

Larval habitat for the avian malaria vector Culex quinquefasciatus (Diptera: Culicidae) in altered mid‐elevation mesic‐dry forests in Hawai'i

Effective management of avian malaria (Plasmodium relictum) in Hawai'i's endemic honeycreepers (Drepanidinae) requires the identification and subsequent reduction or treatment of larval habitat for the mosquito vector, Culex quinquefasciatus (Diptera: Culicidae). We conducted ground surveys, treehole surveys, and helicopter aerial surveys from 2001–2003 to identify all potential larval mosquito habitat within two 100+ ha mesic‐dry forest study sites in Hawai'i Volcanoes National Park, Hawai'i; ‘Ainahou Ranch and Mauna Loa Strip Road. At ‘Ainahou Ranch, anthropogenic sites (43%) were more likely to contain mosquitoes than naturally occurring (8%) sites. Larvae of Cx. quinquefasciatus were predominately found in anthropogenic sites while Aedes albopictus larvae occurred less frequently in both anthropogenic sites and naturally‐occurring sites. Additionally, moderate‐size (≈20–22,000 liters) anthropogenic potential larval habitat had >50% probability of mosquito presence compared to larger‐ and smaller‐volume habitat (<50%). Less than 20% of trees surveyed at ‘Ainahou Ranch had treeholes and few mosquito larvae were detected. Aerial surveys at ‘Ainahou Ranch detected 56% (95% CI: 42–68%) of the potential larval habitat identified in ground surveys. At Mauna Loa Strip Road, Cx. quinquefasciatus larvae were only found in the rock holes of small intermittent stream drainages that made up 20% (5 of 25) of the total potential larval habitat. The volume of the potential larval habitat did not influence the probability of mosquito occurrence at Mauna Loa Strip Road. Our results suggest that Cx. quinquefasciatus abundance, and subsequently avian malaria, may be controlled by larval habitat reduction in the mesic‐dry landscapes of Hawai'i where anthropogenic sources predominate.     

Leaf species identity and combination affect performance and oviposition choice of two container mosquito species

Abstract 1. Resource diversity can be an important determinant of individual and population performance in insects. Fallen parts of plants form the nutritive base for many aquatic systems, including mosquito habitats, but the effect of plant diversity on mosquito production is poorly understood. 2. To determine the effects of diverse plant inputs on larval mosquitoes, experiments were conducted that examined how leaves of Vitis aestivalis, Quercus virginiana, Psychotria nervosa, and Nephrolepis exaltata affected the container species Aedes triseriatus and Aedes albopictus. 3. The hypothesis that leaf species have different effects on larval survival, growth, population performance, and oviposition choice of the two mosquito species was tested. The hypothesis that larval performance of A. albopictus responds additively to combinations of the four plant species was also tested. 4. Larval survival and growth differed among the four leaf species, and oviposition preference differed among the two leaf species examined. Measurements of population performance demonstrated significant variation between leaf treatments. Larval outcomes for A. albopictus were significantly affected by leaf combination, and the hypothesis of additivity could be rejected. 5. These results indicate that individual leaf species are important in determining the performance of container dwelling mosquitoes, which grow larger and survive better on mixed‐species resource than expected, based on an additive model of resource utilisation.     

Nestedness patterns of container-dwelling mosquitoes: effects of larval habitat within variable terrestrial matrices

Distributions of mosquito larvae likely are a consequence of multiple factors, although two commonly studied factors (quality of the larval environment and the terrestrial matrix in which these habitats reside) have rarely and simultaneously been varied in the field to understand distributions of larvae. We monitored aquatic containers of two leaf detritus levels within a forest, prairie, and industrial habitat across five months to understand the temporal and spatial colonization of aquatic invertebrates in Northcentral Illinois, USA. Data were collected monthly on mosquito populations and the composition of other invertebrates colonizing containers. Overall, six species of mosquitoes colonized containers, with Culex restuans and Aedes triseriatus having the highest relative abundances. There were strong seasonal abundance patterns for these two mosquito species, with the dominant species changing over time in the forest habitat. The responses of other mosquito taxa were more variable, with abundances reflective of either the terrestrial matrix or larval habitat quality. High detritus containers supported the highest abundances of most species encountered, regardless of habitat. Non-mosquito taxa were less common numerically, but analyses suggested that some taxa, such as syrphid larvae, often co-occurred with mosquitoes. Nested subset analysis indicated communities were strongly nested, and that both habitat type and detritus level were important in explaining nested patterns of aquatic invertebrates. Our data show that both the larval habitat and the surrounding terrestrial matrix shape patterns of container mosquitoes, and that other container invertebrates vary in similar ways as mosquitoes. Handling editor: K. Martens