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The transmission of Plasmodium within a vertebrate host population is strongly associated with the life history traits of its vector. Therefore the effect of malaria infection on mosquito fecundity and longevity has traditionally received a lot of attention. Several species of malaria parasites reduce mosquito fecundity, nevertheless almost all of the studies have focused only on the first gonotrophic cycle. Yet, during their lifetime, female mosquitoes go through several gonotrophic cycles, which raises the question of whether they are able to compensate the fecundity costs induced by the parasite. The impact of Plasmodium infection on female longevity is not so clear and has produced conflicting results. Here we measured the impact of Plasmodium relictum on its vector's longevity and fecundity during three consecutive gonotrophic cycles. In accordance with previous studies, we observed a negative impact of Plasmodium infection on mosquito (Culex pipiens) fecundity in the first gonotrophic cycle. Interestingly, despite having taken two subsequent uninfected blood meals, the negative impact of malaria parasite persisted. Nevertheless no impact of infection on mosquito longevity was observed. Our results are not in line with the hypothesis that the reduction of fecundity observed in infected mosquitoes is an adaptive strategy of Plasmodium to increase the longevity of its vector. We discuss the different underlying mechanisms that may explain our results.

Sex-biased infections are a recurrent observation in vertebrates. In many species, males are more parasitized than females. Two potentially complementary mechanisms are often suggested to explain this pattern: sexual differences in susceptibility mainly caused by the effect of sex hormones on immunity and differential exposure to parasites. Exposure is mostly a consequence of host behavioural traits, but vector-borne parasitic infections involve another degree of complexity due to the active role of vectors in transmission. Blood-sucking insects may make choices based on cues produced by hosts. Regarding malaria, several studies highlighted a male-biased infection by Plasmodium sp in great tits (Parus major). We hypothesize that the mosquito vector, Culex pipiens, might at least partially cause this bias by being more attracted to male birds. Intrinsic variation associated to bird sex would explain a preference of mosquitoes for males. To test this hypothesis, we provide uninfected mosquitoes with a choice between uninfected male and female nestlings. Mosquito choice is assessed by sex typing of the ingested blood. We did not observe any preference for a given sex. This result does not support our prediction of a preference of mosquitoes for male great tits during the nestling period. In conclusion, mosquitoes do not seem to have an intrinsic preference for male nestlings. However, sexually divergent traits (e.g. behaviour, odour, metabolic rate) present in adults may play a role in the attraction of mosquitoes and should be investigated.

Background In vector-borne diseases, invertebrate hosts are exposed to highly variable quantities of parasites during their blood meal. This heterogeneity may partly explain the overdispersed distribution of parasites within the vector population and the variability in the extrinsic incubation period (EIP) of the parasite. Indeed, the quantity of parasites ingested is often considered as a good predictor of the quantity of parasites that will develop within the vectors, as well as the speed at which they will develop (i.e. EIP). However, density-dependent processes can influence the relationship between parasite burden in the vertebrate host and in vectors, making this relationship unclear at times. Methods Here, we used an avian malaria system to investigate whether the proportion of red blood cells infected by sexual and/or asexual stages of Plasmodium relictum influences the intensity of infection and the EIP within vectors. For this purpose, we experimentally infected 12 birds in order to generate a range of infection intensity. More than 1000 mosquitoes took a blood meal on these hosts, and the development of Plasmodium within the vectors was followed for more than 20 days. Results Our study reveals a negative relationship between the intensity of infection in birds and the time until 10% of mosquitoes become infectious (EIP 10 ). A period of only 4 days was sufficient to detect sporozoites in at least 10% of mosquitoes fed on the most infected hosts. However, the number of sporozoites did not vary significantly according to the vertebrate host intensity of infection, but was positively correlated to the oocyst burden (parasitic stage preceding the sporozoite stage). Conclusions While the quantity of ingested parasites had no impact on oocyst and sporozoite burden in infectious mosquitoes, the EIP 10 was affected. Studies have demonstrated that small changes in the EIP can have a significant effect on the number of mosquitoes living long enough to transmit parasites. Here, we observed a difference of 4–6 days in the detection of the first sporozoites, depending on the intensity of infection of the bitten vertebrate host. Considering that a gonotrophic cycle lasts 3–4 days, the shortened EIP may have significant effects on Plasmodium transmission. Graphical Abstract

Glyphosate is the world's most widely used herbicide. The commercial success of this molecule is due to its nonselectivity and its action, which would supposedly target specific biosynthetic pathways found mainly in plants. Multiple studies have however provided evidence for high sensitivity of many nontarget species to glyphosate and/or to formulations (glyphosate mixed with surfactants). This herbicide, found at significant levels in aquatic systems through surface runoffs, impacts life history traits and immune parameters of several aquatic invertebrates' species, including disease‐vector mosquitoes. Mosquitoes, from hatching to emergence, are exposed to aquatic chemical contaminants. In this study, we first compared the toxicity of pure glyphosate to the toxicity of glyphosate‐based formulations for the main vector of avian malaria in Europe, Culex pipiens mosquito. Then we evaluated, for the first time, how field‐realistic dose of glyphosate interacts with larval nutritional stress to alter mosquito life history traits and susceptibility to avian malaria parasite infection. Our results show that exposure of larvae to field‐realistic doses of glyphosate, pure or in formulation, did not affect larval survival rate, adult size, and female fecundity. One of our two experimental blocks showed, however, that exposure to glyphosate decreased development time and reduced mosquito infection probability by malaria parasite. Interestingly, the effect on malaria infection was lost when the larvae were also subjected to a nutritional stress, probably due to a lower ingestion of glyphosate. Glyphosate is the world's most widely used herbicide. The commercial success of this molecule is due to its nonselectivity and its action, which would supposedly target specific biosynthetic pathways found mainly in plants. However, we have shown that exposure to field‐realistic dose of glyphosate decreases the development time of mosquito larvae and reduces mosquito infection probability by malaria parasite. Interestingly, the effect on malaria infection was lost when the larvae are subjected to a nutritional stress, probably due to a lower ingestion of glyphosate.

Accurate detection and identification of vector-host-parasite systems are key to understanding their evolutionary dynamics and to design effective disease prevention strategies. Traditionally, microscopical and serological techniques were employed to analyse arthropod blood meals for host/parasite detection, but these were limited in taxonomic resolution and only to pre-selected taxa. In recent years, molecular techniques have emerged as a promising alternative, offering enhanced resolution and taxonomic range. While singleplex polymerase chain reaction (PCR) assays were used at first to identify host, vector and parasite components in separate reactions, today multiple primer pairs can be combined in a single reaction, i.e. , multiplex, offering substantial time and cost savings. Nonetheless, despite the potential benefits of multiplex PCR, studies quantifying its efficacy compared to singleplex reactions are scarce. In this study, we used partially digested mosquito blood meals within an avian malaria framework to jointly identify the host, vector and parasite using multiplex DNA metabarcoding, and to compare it with separate singleplex PCRs. We aimed to compare the detection probabilities and taxonomic assignments between both approaches. We found both to have similar performances in terms of detection for the host and the vector, but singleplex clearly outperformed multiplex for the parasite component. We suggest adjusting the relative concentrations of the PCR primers used in the multiplex assay could increase the efficiency of multiplex in detecting all the components of the studied multi-species system. Overall, the results show that multiplex DNA metabarcoding can be an effective approach that could be applied to any vector-borne interaction involving blood-feeding arthropods. Our insights from this proof-of-concept study will help improve laboratory procedures for accurate and cost-efficient medical diagnosis of vector-borne diseases, the spread of which is globally exacerbated by current climate change.

[This corrects the article DOI: 10.1371/journal.pone.0216360.].

Background Changes in host phenotype following parasite infection are often considered as host manipulation when they seem advantageous for the parasite. However, putative cases of host manipulation by parasites are rarely tested in field-realistic conditions. Infection-induced phenotypic change cannot be conclusively considered as host manipulation if no evidence shows that this trait is adaptive for the parasite in the wild. Plasmodium sp., the parasites causing malaria in vertebrates, are hypothesized to “manipulate” their host by making their odour more attractive to mosquitoes, their vector and final host. While this is fairly well supported by studies on mice and humans, studies focusing on avian malaria give contradictory results. Methods In the present study, genotyped birds at different stages (uninfected, acute and chronic) of Plasmodium relictum infection were exposed, in a large outdoor aviary, to their natural vector, the mosquito Culex pipiens . Results After genotyping the blood meals of more than 650 mosquitoes, we found that mosquitoes did not bite infected birds more than they bit them before infection, nor more than they bit uninfected hosts. Conclusions Our study highlights the importance of testing ecological behaviours under natural conditions and suggests that different processes might be at play in mammals and birds regarding potential manipulation of attractiveness by malaria parasites. Graphical Abstract

Background Parasites are able to alter numerous aspects of their hosts’ life history, behaviour and distribution. One central question in parasitology is to determine the degree of impact that parasites have on their hosts. Laboulbeniales (Fungi: Ascomycota) are ectoparasitic fungi of arthropods. Even though these fungi are widely distributed, little is known about their ecology and their possible physiological effects on their hosts. We used a highly specific bat fly-fungi association to assess the effect of these fungal parasites on their dipteran hosts. Methods We collected bat flies (Diptera: Nycteribiidae) belonging to two species, Nycteribia schmidlii and Penicillidia conspicua from their bat host Miniopterus schreibersii (Chiroptera: Miniopteridae). We experimentally tested the effect of infection on the lifespan of bat flies. Results The prevalence of Laboulbeniales fungi was 17.9% in N. schmidlii and 64.8% in P. conspicua . Two fungi species were identified, Arthrorhynchus eucampsipodae and A. nycteribiae , both showing strict host specificity with N. schmidlii and P. conspicua , respectively. We found that fungal infection reduced by half the survival rate of P. conspicua regardless of sex, whereas N. schmidlii was not affected by the infection. Moreover, the intensity of infection showed negative correlation with the lifespan of P. conspicua . Conclusions To our knowledge, this is the first indication that fungal infection can alter bat fly survival and thus may play a significant role in the population dynamics of these bat ectoparasites.

Malaria, a vector-borne disease caused by Plasmodium spp ., remains a major global cause of mortality. Optimization of disease control strategies requires a thorough understanding of the processes underlying parasite transmission. While the number of transmissible stages (gametocytes) of Plasmodium in blood is frequently used as an indicator of host-to-mosquito transmission potential, this relationship is not always clear. Significant effort has been made in developing molecular tools that improve gametocyte density estimation and therefore prediction of mosquito infection rates. However a significant level of uncertainty around estimates remains. The weakness in the relationship between gametocyte burden, measured from a blood sample, and the mosquito infection rate could be explained by a non-homogeneous distribution of gametocytes in the bloodstream. The estimated gametocyte density would then only be a single snapshot that does not reflect the host infectivity. This aspect of Plasmodium infection, however, remains largely neglected. In both humans and birds, we found here that the gametocyte densities differed depending on which side of the body the sample was taken, suggesting that gametocytes are not homogeneously distributed within the vertebrate host. We observed a fluctuating asymmetry, in other words, the extremity of the body with the highest density of parasites is not always the same from one individual to another. An estimation of gametocyte density from only one blood sample, as is commonly measured, could, therefore, over- or underestimated the infectivity of gametocyte carriers. This might have important consequences on the epidemiology of the disease since we show that this variation influences host-to-mosquito transmission. Vectors fed on the least infected body part had a lower parasite burden than those fed on the most infected part. The heterogeneous distribution of gametocytes in bloodstream should be considered to improve diagnosis and test new malaria control strategies.

Vector mosquitoes contribute significantly to the global burden of diseases in humans, livestock and wildlife. As such, the spatial distribution and abundance of mosquito species and their surveillance cannot be ignored. Here, we surveyed mosquito species across major tourism hotspots in semi-arid Botswana, including, for the first time, the Central Kalahari Game Reserve. Our results reported several mosquito species across seven genera, belonging to Aedes, Anopheles, Culex, Mansonia, Mimomyia, Coquillettidia and Uranotaenia. These results document a significant species inventory that may inform early warning vector-borne disease control systems and likely help manage the risk of emerging and re-emerging mosquito-borne infections.