Cannibalism and predation among larvae of the Anopheles gambiae complex

Among the aquatic developmental stages of the Anopheles gambiae complex (Diptera: Culicidae), both inter- and intra-specific interactions influence the resulting densities of adult mosquito populations. For three members of the complex, An. arabiensis Patton, An. quadriannulatus (Theobald) and An. gambiae Giles sensu stricto, we investigated some aspects of this competition under laboratory conditions. First-instar larvae were consumed by fourth-instar larvae of the same species (cannibalism) and by fourth-instar larvae of other sibling species (predation). Even when larvae were not consumed, the presence of one fourth-instar larva caused a significant reduction in development rate of first-instar larvae. Possible implications of these effects for population dynamics of these malaria vector mosquitoes are discussed.     

Anopheles arabiensis and An. quadriannulatus resistance to DDT in South Africa

The malaria control programme of KwaZulu‐Natal Province, South Africa, includes Mamfene and Mlambo communities. Western‐type houses there are currently sprayed with deltamethrin, whereas traditional houses are sprayed with DDT for malaria control. In 2002, mosquitoes of the Anopheles gambiae complex (Diptera: Culicidae) were collected from DDT‐sprayed houses, by window exit traps, and from man‐baited nets outdoors. Larval collections were also carried out at Mzinweni Pan near Mlambo. Species of the An. gambiae complex were identified by rDNA polymerase chain reaction assay. The majority of samples collected by window trap and baited nets were identified as the malaria vector An. arabiensis Patton, with a few An. merus Dönitz and An. quadriannulatus (Theobald). The larval collections were predominantly An. quadriannulatus with a small number of An. arabiensis. Standard WHO insecticide susceptibility tests using 4% DDT and 0.05% deltamethrin were performed on both wild‐caught females and laboratory‐reared progeny from wild‐caught females. Wild‐caught An. arabiensis samples from window traps gave 63% and 100% mortality 24‐h post‐exposure to DDT or deltamethrin, respectively. Wild‐caught An. arabiensis samples from man‐baited net traps gave 81% mortality 24‐h post‐exposure to DDT. The F₁ progeny from 22 An. arabiensis females showed average mortality of 86.5% 24‐h post‐exposure to DDT. Less than 80% mortality was recorded from five of these families. Biochemical analyses of samples from each of the families revealed comparatively high levels of glutathione‐S‐transferases and non‐specific esterases in some families, but without significant correlation to bioassay results. Wild‐caught An. quadriannulatus larvae were reared through to adults and assayed on 4% DDT, giving 47% (n = 36) mortality 24‐h post‐exposure. Finding DDT resistance in the vector An. arabiensis, close to the area where we previously reported pyrethroid‐resistance in the vector An. funestus Giles, indicates an urgent need to develop a strategy of insecticide resistance management for the malaria control programmes of southern Africa.     

Larvae of cryptic species of Anopheles gambiae respond differently to cues of predation risk

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Conspecific Sharing of Breeding Sites by Anopheline Female Mosquitoes (Diptera: Culicidae) Inferred from Microsatellite Markers

The number of Anopheles gambiae and Anopheles arabiensis females that used each of the 33 sampled breeding sites in west Kenya was estimated by microsatellite markers and related statistics to test the hypothesis that conspecific females share aquatic sites. Totally, 166 An. gambiae and 168 An. arabiensis larvae were identified and were genotyped. The mean number of larvae per breeding site was 8.3 for An. gambiae and 8.4 for An. arabiensis. The likelihood method estimated that, for An. gambiae, the mean number of females that would have laid eggs per breeding site was 5.2 and ranged from 2 to 9, and for An. arabiensis, the mean was 5.0 with a range of 2–10. The clustering method estimated that the mean number of females laying eggs per breeding site was 6.8 for An. gambiae. The results provide molecular evidence that females of one or both species share breeding sites.     

Insecticide resistance monitoring of field‐collected Anopheles gambiae s.l. populations from Jinja,eastern Uganda,identifies high levels of pyrethroid resistance

Insecticide resistance in the malaria vector Anopheles gambiae s.l. (Diptera: Culicidae) threatens insecticide‐based control efforts, necessitating regular monitoring. We assessed resistance in field‐collected An. gambiae s.l. from Jinja, Uganda using World Health Organization (WHO) biosassays. Only An. gambiae s.s. and An. arabiensis (?70%) were present. Female An. gambiae exhibited extremely high pyrethroid resistance (permethrin LT₅₀ > 2 h; deltamethrin LT₅₀ > 5 h). Female An. arabiensis were resistant to permethrin and exhibited reduced susceptibility to deltamethrin. However, while An. gambiae were DDT resistant, An. arabiensis were fully susceptible. Both species were fully susceptible to bendiocarb and fenitrothion. Kdr 1014S has increased rapidly in the Jinja population of An. gambiae s.s. and now approaches fixation (?95%), consistent with insecticide‐mediated selection, but is currently at a low frequency in An. arabiensis (0.07%). Kdr 1014F was also at a low frequency in An. gambiae. These frequencies preclude adequately‐powered tests for an association with phenotypic resistance. PBO synergist bioassays resulted in near complete recovery of pyrethroid susceptibility suggesting involvement of CYP450s in resistance. A small number (0.22%) of An. gambiae s.s. ×An. arabiensis hybrids were found, suggesting the possibility of introgression of resistance alleles between species. The high levels of pyrethroid resistance encountered in Jinja threaten to reduce the efficacy of vector control programmes which rely on pyrethroid‐impregnated bednets or indoor spraying of pyrethroids.     

Dynamics of cannibalism in equal‐aged cohorts of Spodoptera frugiperda

1. Antagonistic interactions in herbivorous insects are often density‐dependent, so rates are predicted to vary dynamically over time as density changes. Fatal intraspecific interactions, especially cannibalism, occur between equal‐aged larvae in young first‐ and second‐instar Spodoptera frugiperda (J.E. Smith). 2. A cannibalism experiment was conducted, starting with seven different densities of neonate S. frugiperda larvae, each replicated 50 times. Larvae were examined daily for the duration of the first and second instars (7 days). Seven‐day mortality was density‐dependent. 3. A stochastic mathematical model was developed in which per‐capita mortality from antagonistic interactions among equal‐aged larvae varies dynamically as density changes. A maximum likelihood method was developed to estimate the conditional per‐capita mortality rate from antagonistic interactions given an intraspecific encounter. An alternative model with mean‐mortality from antagonistic interactions that depends only on the initial larval density was also developed. 4. The models were fitted to the experimental data, and compared using log‐likelihood. The dynamic model fitted the cannibalism data significantly better than the time‐averaged mortality model for all starting densities for the experimental data, implying that density‐dependent mortality varied dynamically over time even within short 7‐day periods. 5. The conditional per‐capita mortality rate from antagonistic interactions was also density‐dependent, possibly because encounters became more aggregated at higher density, or because the probability that a larva died from the interaction was higher at higher density, or both.     

Distribution and Frequency of kdr Mutations within Anopheles gambiae s.l. Populations and First Report of the Ace.1G119S Mutation in Anopheles arabiensis from Burkina Faso (West Africa)

An entomological survey was carried out at 15 sites dispersed throughout the three eco-climatic regions of Burkina Faso (West Africa) in order to assess the current distribution and frequency of mutations that confer resistance to insecticides in An. gambiae s.l. populations in the country. Both knockdown (kdr) resistance mutation variants (L1014F and L1014S), that confer resistance to pyrethroid insecticides, were identified concomitant with the ace-1 G119S mutation confirming the presence of multiple resistance mechanisms in the An. gambiae complex in Burkina Faso. Compared to the last survey, the frequency of the L1014F kdr mutation appears to have remained largely stable and relatively high in all species. In contrast, the distribution and frequency of the L1014S mutation has increased significantly in An. gambiae s.l. across much of the country. Furthermore we report, for the first time, the identification of the ace.1 G116S mutation in An. arabiensis populations collected at 8 sites. This mutation, which confers resistance to organophosphate and carbamate insecticides, has been reported previously only in the An. gambiae S and M molecular forms. This finding is significant as organophosphates and carbamates are used in indoor residual sprays (IRS) to control malaria vectors as complementary strategies to the use of pyrethroid impregnated bednets. The occurrence of the three target-site resistance mutations in both An. gambiae molecular forms and now An. arabiensis has significant implications for the control of malaria vector populations in Burkina Faso and for resistance management strategies based on the rotation of insecticides with different modes of action.     

Species Shifts in the Anopheles gambiae Complex: Do LLINs Successfully Control Anopheles arabiensis?

Introduction

High coverage of conventional and long-lasting insecticide treated nets (ITNs and LLINs) in parts of E Africa are associated with reductions in local malaria burdens. Shifts in malaria vector species ratio have coincided with the scale-up suggesting that some species are being controlled by ITNs/LLINs better than others.

Methods

Between 2005–2006 six experimental hut trials of ITNs and LLINs were conducted in parallel at two field stations in northeastern Tanzania; the first station was in Lower Moshi Rice Irrigation Zone, an area where An. arabiensis predominates, and the second was in coastal Muheza, where An. gambiae and An. funestus predominate. Five pyrethroids and one carbamate insecticide were evaluated on nets in terms of insecticide-induced mortality, blood-feeding inhibition and exiting rates.

Results

In the experimental hut trials mortality of An. arabiensis was consistently lower than that of An. gambiae and An. funestus. The mortality rates in trials with pyrethroid-treated nets ranged from 25–52% for An. arabiensis, 63–88% for An. gambiae s.s. and 53–78% for An. funestus. All pyrethroid-treated nets provided considerable protection for the occupants, despite being deliberately holed, with blood-feeding inhibition (percentage reduction in biting rates) being consistent between species. Veranda exiting rates did not differ between species. Percentage mortality of mosquitoes tested in cone bioassays on netting was similar for An. gambiae and An. arabiensis.

Conclusions

LLINs and ITNs treated with pyrethroids were more effective at killing An. gambiae and An. funestus than An. arabiensis. This could be a major contributing factor to the species shifts observed in East Africa following scale up of LLINs. With continued expansion of LLIN coverage in Africa An. arabiensis is likely to remain responsible for residual malaria transmission, and species shifts might be reported over larger areas. Supplementary control measures to LLINs may be necessary to control this vector species.     

Variation in host preferences of malaria mosquitoes is mediated by skin bacterial volatiles

The host preferences of the anthropophilic mosquito species in the Anopheles gambiae complex (Diptera: Culicidae) are mediated by skin bacterial volatiles. However, it is not known whether these mosquitoes respond differentially to skin bacterial volatiles from non‐human host species. In this study, the responses of two malaria mosquito species in the An. gambiae complex, Anopheles gambiae s.s. (hereafter, An. gambiae) and Anopheles arabiensis, with different host preferences, to volatiles released from skin bacteria were tested. Skin bacteria collected from human, cow and chicken skin significantly increased trap catches; traps containing bacteria collected from human skin caught the highest proportions of An. gambiae and An. arabiensis. Traps with bacteria of human origin caught a significantly higher proportion of An. gambiae than of An. arabiensis, whereas bacterial volatiles from the chicken attracted significantly higher numbers of An. arabiensis than of An. gambiae. Additionally, An. gambiae showed a specialized response to volatiles from four specific bacteria, whereas An. arabiensis responded equally to all species of bacteria tested. Skin bacterial volatiles may therefore play important roles in guiding mosquitoes with different host preferences. The identification of these bacterial volatiles can contribute to the development of an odour blend that attracts mosquitoes with different host preferences.     

Shift in species composition in the Anopheles gambiae complex after implementation of long‐lasting insecticidal nets in Dielmo,Senegal

Long‐lasting insecticidal nets (LLINs) and indoor residual spraying (IRS) are the cornerstones of malaria vector control. However, the effectiveness of these control tools depends on vector ecology and behaviour, which also largely determine the efficacy of certain Anopheles mosquitoes (Diptera: Culicidae) as vectors. Malaria vectors in sub‐Saharan Africa are primarily species of the Anopheles gambiae complex, which present intraspecific differences in behaviour that affect how they respond to vector control tools. The focus of this study is the change in species composition in the An. gambiae complex after the implementation of LLINs in Dielmo, Senegal. The main findings referred to dramatic decreases in the proportions of Anopheles coluzzii and An. gambiae after the introduction of LLINs, and an increase in the proportion of Anopheles arabiensis. Two years after LLINs were first introduced, An. arabiensis remained the most prevalent species and An. gambiae had begun to rebound. This indicated a need to develop additional vector control tools that can target the full range of malaria vectors.     

Modification of Anopheles gambiae distribution at high altitudes in Madagascar

In Madagascar, Anopheles gambiae has been found below altitudes of 1,000 m. We sampled An. gambiae sensu lato (sl) between 2008 and 2010 in the Central Highlands of Madagascar at altitudes over 1,200 m. The study site consists of rainforest, rainforest edge, and an open savanna biotope. Anopheles gambiae and An. arabiensis, as well as molecular forms of An. gambiae, were identified molecularly. An. gambiae accounted for 26.7% at the edge of the rainforest and 2.3% in the open savanna biotope. One specimen of this species was caught in the forest. An. arabiensis accounted for 66.3% at the edge of the rainforest and 97.7 % in the open savanna biotope. All An. gambiae adults tested belonged to the S molecular form. An. gambiae is present at high altitudes in Madagascar, with a high prevalence at the rainforest edge. Several factors, including the appearance of new favorable biotopes, recolonization after a reduction of indoor vector control, and climate change, may contribute to its distribution. The changing distribution of An. gambiae may have consequences for the distribution and incidence of malaria in the Malagasy Highlands.     

The impact of host species and vector control measures on the fitness of African malaria vectors

Many malaria vector mosquitoes in Africa have an extreme preference for feeding on humans. This specialization allows them to sustain much higher levels of transmission than elsewhere, but there is little understanding of the evolutionary forces that drive this behaviour. In Tanzania, we used a semi-field system to test whether the well-documented preferences of the vectors, Anopheles arabiensis and Anopheles gambiae sensu stricto (s.s.) for cattle and humans, respectively, are predicted by the fitness they obtain from host-seeking on these species relative to other available hosts. Mosquito fitness was contrasted, when humans were fully exposed and when they were protected by a typical bednet. The fitness of both vectors varied between host species. The predicted relationship between host preference and fitness was confirmed in An. arabiensis, but not in An. gambiae s.s., whose fitness was similar on humans and other mammals. Use of typical, imperfect bednets generated only minor reductions in An. gambiae s.s. feeding success and fitness on humans, but was predicted to generate a significant reduction in the lifetime reproductive success of An. arabiensis on humans relative to cows. This supports the hypothesis that such human-protective measures could additionally benefit malaria control by increasing selection for zoophily in vectors.     

Islands and Stepping-Stones: Comparative Population Structure of Anopheles gambiae sensu stricto and Anopheles arabiensis in Tanzania and Implications for the Spread of Insecticide Resistance

Population genetic structures of the two major malaria vectors Anopheles gambiae s.s. and An. arabiensis, differ markedly across Sub-Saharan Africa, which could reflect differences in historical demographies or in contemporary gene flow. Elucidation of the degree and cause of population structure is important for predicting the spread of genetic traits such as insecticide resistance genes or artificially engineered genes. Here the population genetics of An. gambiae s.s. and An. arabiensis in the central, eastern and island regions of Tanzania were compared. Microsatellite markers were screened in 33 collections of female An. gambiae s.l., originating from 22 geographical locations, four of which were sampled in two or three years between 2008 and 2010. An. gambiae were sampled from six sites, An. arabiensis from 14 sites, and both species from two sites, with an additional colonised insectary sample of each species. Frequencies of the knock-down resistance (kdr) alleles 1014S and 1014F were also determined. An. gambiae exhibited relatively high genetic differentiation (average pairwise F_ST = 0.131), significant even between nearby samples, but without clear geographical patterning. In contrast, An. arabiensis exhibited limited differentiation (average F_ST = 0.015), but strong isolation-by-distance (Mantel test r = 0.46, p = 0.0008). Most time-series samples of An. arabiensis were homogeneous, suggesting general temporal stability of the genetic structure. An. gambiae populations from Dar es Salaam and Bagamoyo were found to have high frequencies of kdr 1014S (around 70%), with almost 50% homozygote but was at much lower frequency on Unguja Island, with no. An. gambiae population genetic differentiation was consistent with an island model of genetic structuring with highly restricted gene flow, contrary to An. arabiensis which was consistent with a stepping-stone model of extensive, but geographically-restricted gene flow.     

Relationship of larval desiccation to Anopheles gambiae Giles and An. arabiensis Patton survival

The relationship between mosquito 4^th instar larval desiccation and survival to adulthood was explored by three methods in the laboratory. Two colonies of Anopheles arabiensis and one of Anopheles gambiae were studied. We found significant differences in tolerance to desiccation among all three stocks suggesting an intra‐ and interspecific genetic component to desiccation tolerance. An. arabiensis KGB, originating from Zimbabwe about 1975, had a much‐reduced desiccation tolerance compared to An. gambiae G3, colonized in the Gambia in 1975, and An. arabiensis DONGOLA which originated in Sudan in 2004. Individuals of the G3 stock survived desiccation of times up to 40 min with survival of 0.52. The degree of difference in tolerance between G3 and DONGOLA was smallest and was detected by one of three experimental methods. Mass losses of individuals that were weighed individually and survived to adulthood averaged 27% and 29% for G3 and DONGOLA and 20% for the less tolerant KGB stock, respectively. Such differences in survival in transiently dry larval habitats may account in part for differences in the distribution of these species and karyotypes.     

The association between the phytoplankton, Rhopalosolen species (Chlorophyta; Chlorophyceae), and Anopheles gambiae sensu lato (Diptera: Culicidae) larval abundance in western Kenya

Algae are important food resources of the larvae of the African malaria vectors, Anopheles gambiae Giles and Anopheles arabiensis Patton (Anopheles gambiae sensu lato), and other zooplankton, but empirical evidence remains meager about the agal flora in ephemeral water bodies. The animals present in natural aquatic habitats in western Kenya were sampled from July to November 2002 to study abiotic and biotic environmental factors determining A. gambiae sl larval abundance. The five highest concentrations of third and fourth instars and pupae (hereafter referred to as old-stage larvae) were sampled in conjunction with the unicellular epizoic green algae, Rhopalosolen species (Chlorophyta; Chlorophyceae). Canonical correspondence analysis revealed that the presence of Rhopalosolen species was the most important determinant of the animal assemblage. The density of old-stage A. gambiae sl larvae was positively correlated with the presence of Rhopalosolen species, but the density of first and second instars of A. gambiae sl was not. The water bodies with Rhopalosolen sp. yielded larger mosquitoes in spite of the higher density of larvae. We demonstrated that the productivity of water bodies in terms of the larvae of malaria vectors can differ in magnitude depending on the agal flora. We discuss phytoplankton as a regulator of mosquito larval populations.     

Interference and competitive behaviour of the aphid predators,Chrysoperla carnea andCoccinella septempunctata in the laboratory

Interference and competition betweenChrysoperla carnea (Stephens) andCoccinella septempunctata L. was investigated in the absence and presence of aphid prey. When larvae of similar vigour encountered each other,C. carnea larvae were superior toC. septempunctata. Otherwise the larger of 2 individuals always killed the smaller. In the absence of preyC. carnea adults were attacked by their own 2nd and 3rd instar larvae as well as by 3rd instar larvae ofC. septempunctata. In all encounters the adults ofC. septempunctata were superior. Since the eggs ofC. carnea are protected to some extent by being on egg stalks, they were less susceptible to cannibalism and predation thanC. septempunctata eggs which are deposited in batches directly on plant leaves. In the presence of prey cannibalism and predation were reduced, especially between larvae. Only eggs and 1st instar larvae were endangered. In the present experimentsC. carnea showed a slight superiority overc. septempunctata. Paper presented at the 17th International Congress of Entomology held in August 20–26 1984 in Hamburg, F.R. Germany.     

Effects of alternative food on cannibalism and herbivore suppression by carabid larvae

1. Predator and alternative food density are important factors influencing herbivore suppression by generalist predators. Herbivore suppression can be reduced if predators forage preferentially on alternative foods. Cannibalism can increase at high predator densities, further reducing herbivore suppression. However, complex interactions are possible, as alternative food can increase predator abundance and survival restoring top‐down effects on herbivores. 2. In two species of carabid larvae (Poecilus chalcites and Anisodactylus ovularis), we studied how alternative foods (fly pupae and grass seeds) and predator density affect predation of black cutworm larvae and how alternative foods affect cannibalism among carabid larvae. 3. Adding alternative food to microcosms generally reduced total predation of cutworms. However, the strength of this effect was dependent on carabid species, larval density, and food type. 4. Increasing larval density from one to three per microcosm reduced per‐capita predation by both species irrespective of alternative food treatment. 5. Alternative food reduced cannibalism in both carabid species and increased survival of carabid larvae in field plots, such that twice as many were captured in plots subsidised with pupae than plots with no alternative food. 6. These results provide new insight into the complex interactions that influence predator survival and herbivore suppression in resource diverse habitats by demonstrating the primacy of intraguild interactions among carabid larvae.     

DNA probes for species identification of mosquitoes in the Anopheles gambiae complex

Abstract. Identification of species within the Anopheles gambiae Giles species complex is essential for the correct evaluation of malaria vector ecology studies and control programmes. The development of DNA probes to distinguish species of the An.gambiae complex is described. Genomic libraries were prepared for four members of the An.gambiae complex. These were screened using radiolabeled DNA from different species of An. gambiae sensu lato and a number of clones selected on the basis of their species specificity. These clones could be divided into two groups, each containing homologous sequences. Sequences homologous to group 1 inserts are highly reiterated in the genomes of Anopheles arabiensis Patton and Anopheles merus Dönitz, present in low copy number in Anopheles melas Theobald, but were not detected in Anopheles gambiae sensu stricto. Studies on the organization of this sequence in the genome of An.arabiensis show that homologous sequences are male specific and interspersed within the chromatin. Sequences homologous to group 2 inserts are highly repeated in the genomes of An.merus and An.melas, but present in low copy number in An.gambiae s.s. and An.arabiensis. Group 2 homologous sequences are not sex-specific in the species tested and appear to be tandemly repeated. When used as hybridization probes, these sequences provide a sensitive means for the identification of species within the Anopheles gambiae complex.     

Life history consequences of larval foraging depth differ between two competing Anopheles mosquitoes

1. Anopheline larvae are surface feeders and allocate most of their time to search for food at the water surface. However, species of the Anopheles gambiae Giles complex may also show bottom feeding. The consequences of this foraging tactic for life history are unknown, yet may be relevant to understand inter‐specific competition patterns. 2. The diving ability and activity of larvae of the main African malaria vectors, An. coluzzii and An. gambiae, at two different water depths (14 and 30 cm) were assessed. We further explored the biological relevance of diving for food harvesting by monitoring key life history traits in two species treatments (single or mixed species) and two food treatments (surface or bottom feeding). 3. Overall, An. coluzzii larvae showed more diving activity than An. gambiae. When feeding at the bottom both species, and especially An. gambiae, showed a delayed emergence and a reduced emergence rate. Moreover, An. gambiae also suffered a reduced wing length. 4. Mixed‐species rearing had a detrimental effect on the life history traits of An. gambiae but not on An. coluzzii, suggesting a competitive advantage for the latter in our experimental conditions. 5. The present results confirm that anopheline larvae are able to forage for food at the bottom of their breeding site and that An. coluzzii shows a superior diving activity than An. gambiae and this at a lower cost. These behavioural differences probably reflect specific adaptations to different aquatic habitats, and may be important in shaping species distributions and the population biology of these important vector mosquitoes.     

Larvicidal efficacy ofBacillus thuringiensis var. israelensis andBacillus sphaericus onAnopheles arabiensis in Ethiopia

The second instar larvae of the malaria vector mosquito,Anopheles arabiensis,were more susceptible to Bacillus thuringiensis var. israelensis (IPS-82) and B. sphaericus (SPH-88) than the third instar larvae. The LC ₅₀ values were 1.0 Μg^I-1 and 1.8 Μg_I-1 for IPS-82 against second and third instar larvae respectively, after 48 h of exposure. The LC₅₀ values for SPH-88 were 3.6 Μg {siI-1} against the second instar larvae and 7.6 Μg_I-1 against the third instar larvae of An. arabiensis. The larvicidal efficacy of SPH-88 was significantly less than IPS-82. The potential of IPS-82 for the control of An. arabiensis in malaria endemic areas is promising.