Male-killing Bacteria Research Articles

Male-killer symbiont screening reveals novel associations in Adalia ladybirds.

While male-killing bacteria are known to infect across arthropods, ladybird beetles represent a hotspot for these symbioses. In some host species, there are multiple different symbionts that vary in presence and frequency between populations. To further our understanding of spatial and frequency variation, we tested for the presence of three male-killing bacteria: Wolbachia , Rickettsia and Spiroplasma , in two Adalia ladybird species from a previously unexplored UK population. The two-spot ladybird, A. bipunctata, is known to harbour all three male-killers, and we identified Spiroplasma infection in the Merseyside population for the first time. However, in contrast to previous studies on two-spot ladybirds from continental Europe, evidence from egg-hatch rates indicates the Spiroplasma strain present in the Merseyside population does not cause embryonic male-killing. In the related ten-spot ladybird, A. decempunctata, there is only one previous record of a male-killing symbiont, a Rickettsia , which we did not detect in the Merseyside sample. However, PCR assays indicated the presence of a Spiroplasma in a single A. decempunctata specimen. Marker sequence indicated that this Spiroplasma was divergent from that found in sympatric A. bipunctata. Genome sequencing of the Spiroplasma -infected A. decempunctata additionally revealed the presence of cobionts in the form of a Centistes parasitoid wasp and the parasitic fungi Beauveria. Further study of A. decempunctata from this population is needed to resolve whether it is the ladybird or wasp cobiont that harbours Spiroplasma , and to establish the phenotype of this strain. These data indicate first that microbial symbiont phenotype should not be assumed from past studies conducted in different locations, and second that cobiont presence may confound screening studies aimed to detect the frequency of a symbiont in field collected material from a focal host species.

The son-killer microbe Arsenophonus nasoniae is a widespread associate of the parasitic wasp Nasonia vitripennis in Europe

Heritable microbes that exhibit reproductive parasitism are common in insects. One class of these are the male-killing bacteria, which are found in a broad range of insect hosts. Commonly, our knowledge of the incidence of these microbes is based on one or a few sampling sites, and the degree and causes of spatial variation are unclear. In this paper, we examine the incidence of the son-killer microbe Arsenophonus nasoniae across European populations of its wasp host, Nasonia vitripennis. In preliminary work, we noticed two female N. vitripennis producing highly female biased sex ratios in a field study from the Netherlands and Germany. When tested, the brood from Germany was revealed to be infected with A. nasoniae. We then completed a broad survey in 2012, in which fly pupal hosts of N. vitripennis were collected from vacated birds’ nests from four European populations, N. vitripennis wasps allowed to emerge and then tested for A. nasoniae presence through PCR assay. We then developed a new screening methodology based on direct PCR assays of fly pupae and applied this to ethanol-preserved material collected from great tit (Parus major) nests in Portugal. These data show A. nasoniae is found widely in European N. vitripennis, being present in Germany, the UK, Finland, Switzerland and Portugal. Samples varied in the frequency with which they carry A. nasoniae, from being rare to being present in 50% of the pupae parasitised by N. vitripennis. Direct screening of ethanol-preserved fly pupae was an effective method for revealing both wasp and A. nasoniae infection, and will facilitate sample transport across national boundaries. Future research should examine the causes of variation in frequency, in particular testing the hypothesis that N. vitripennis superparasitism rates drive the variation in A. nasoniae frequency through providing opportunities for infectious transmission.

The impact of intrinsic and extrinsic factors on the epidemiology of male‐killing bacteria

Many arthropod species are vulnerable to infection by bacteria that kill the male offspring of their female hosts. These male‐killing bacteria (MKB) can greatly impact the ecology and evolution of their hosts, particularly when MKB prevalence is high and persistent through time. However, we still do not have a holistic understanding of the key determinants of MKB ecological epidemiology, particularly regarding the interplay between host‐MKB traits and environmental conditions. Here, we derived a mathematical model to analyse the role that several intrinsic and extrinsic factors play in determining MKB epidemiology, and how these factors interact with one another. We found that the invasion threshold and prevalence of MKB is determined by transmission rate and the rate at which female hosts recover from MKB infections. The invasion and prevalence of MKB is also highly sensitive to the extent that MKB can reduce intra‐brood competition by killing male offspring. Environmentally‐induced periodic changes to the epidemiological characteristics of MKB caused a sharp decline and a slow recovery of MKB prevalence and, in some cases, environmental disturbance can drive MKB extinct. Furthermore, the magnitude of the impact that environmental disturbance had on the dynamics of MKB prevalence was heavily modulated by intrinsic factors, particularly intra‐brood competition. This is the first study to explore how both intrinsic and extrinsic factors interact to influence the dynamics of MKB infections over large timescales; our findings are central to predicting the current and future impacts of MKB on host populations.

Sex determination systems as the interface between male-killing bacteria and their hosts.

Arthropods host a range of sex-ratio-distorting selfish elements, including diverse maternally inherited endosymbionts that solely kill infected males. Male-killing heritable microbes are common, reach high frequency, but until recently have been poorly understood in terms of the host–microbe interaction. Additionally, while male killing should generate strong selection for host resistance, evidence of this has been scant. The interface of the microbe with host sex determination is integral to the understanding of how death is sex limited and how hosts can evolve evasion of male killing. We first review current knowledge of the mechanisms diverse endosymbionts use to induce male-specific death. We then examine recent evidence that these agents do produce intense selection for host nuclear suppressor elements. We argue, from our understanding of male-killing mechanisms, that suppression will commonly involve evolution of the host sex determination pathways and that the host's response to male-killing microbes thus represents an unrecognized driver of the diversity of arthropod sex determination. Further work is required to identify the genes and mechanisms responsible for male-killing suppression, which will both determine the components of sex determination (or other) systems associated with suppressor evolution, and allow insight into the mechanism of male killing itself.

Metagenomic survey of bacteria associated with the invasive ladybird Harmonia axyridis (Coleoptera: Coccinellidae)

The Asian ladybird Harmonia axyridis is an invasive insect in Europe and the Americas and is a great threat to the environment in invaded areas. The situation is exacerbated by the fact that non native species are resistant to many groups of parasites that attack native insects. However, very little is known about the complex microbial community associated with this insect. This study based on sequencing 16S rRNA genes in extracted metagenomic DNA is the first research on the bacterial flora associated with H. axyridis. Lady beetles were collected during hibernation from wind turbines in Poland. A mean ± SD of 114 ± 35 species of bacteria were identified. The dominant phyla of bacteria recorded associated with H. axyridis were Actinobacteria, Proteobacteria, Firmicutes and Bacteroidetes. Representatives of these phyla are common in the environment, e.g. in the soil, and are often identified as the dominant bacteria associated with arthropods. We also identified animal pathogenic bacteria, such as Burkholderia, Rhodococcus, Chlamydiae and Anaplasmataceae spp. (Neorickettsia helminthoeca and Ehrlichia ovina). We also identified Wolbachia pipientis in a single beetle. This bacterium is a causative agent of reproductive alterations in arthropods. These results support the enemy release hypothesis in the case of this ladybird invasion. Pathogenic bacteria were recorded in only a few samples. Moreover, male-killing bacteria such as Spiroplasma spp., Wolbachia spp. and Rickettsia spp. were only recorded in single insects so they cannot be responsible for the observed alterations in the sex-ratio of the ladybird population studied.

Variation of clutch size and trophic egg proportion in a ladybird with and without male-killing bacterial infection

Maternally inherited bacterial endosymbionts can kill male embryos of their arthropod hosts to enhance the transmission efficiency of the endosymbionts. The resources from killed male eggs can be reallocated to infected female hatchlings as additional maternal investment. As a result, the number of offspring per patch and the maternal investment per offspring are expected to differ from the original optimal values for the host mother. Thus, in response to infection, these trait values should be adjusted to maximize the lifetime reproductive success of host females and the fitness of inherited endosymbionts as well. Here, we examined clutch size, egg size, and the proportion of trophic eggs (i.e., production of unhatched eggs, a maternal phenotype) per clutch of host mothers infected with male-killing bacteria. First, we developed a mathematical model to predict the optimal clutch size and trophic egg proportion in uninfected and infected females. Next, we experimentally compared these life-history traits in a ladybird, Harmonia yedoensis, between females infected or uninfected with male-killing Spiroplasma bacteria. Consistent with our predictions, clutch size was larger, egg size was smaller, and trophic egg proportion was lower in infected H. yedoensis females, compared with uninfected females. To our knowledge, this is the first empirical demonstration of variation in these life-history traits depending on infection with bacterial endosymbionts.

Evolution of early male-killing in horizontally transmitted parasites.

Early male-killing (MK) bacteria are vertically transmitted reproductive parasites which kill male offspring that inherit them. Whereas their incidence is well documented, characteristics allowing originally non-MK bacteria to gradually evolve MK ability remain unclear. We show that horizontal transmission is a mechanism enabling vertically transmitted bacteria to evolve fully efficient MK under a wide range of host and parasite characteristics, especially when the efficacy of vertical transmission is high. We also show that an almost 100% vertically transmitted and 100% effective male-killer may evolve from a purely horizontally transmitted non-MK ancestor, and that a 100% efficient male-killer can form a stable coexistence only with a non-MK bacterial strain. Our findings are in line with the empirical evidence on current MK bacteria, explain their high efficacy in killing infected male embryos and their variability within and across insect taxa, and suggest that they may have evolved independently in phylogenetically distinct species.

Assessing the impact of male-killing bacteria on the spread of a sexually transmitted infection

Theory predicts that the epidemiology of sexually transmitted infections (STIs) will be affected by any factor that alters the mating biology of its host. In insects, the presence of sex ratio-distorting symbionts may modulate population sex ratio, altering male and female encounter rates and creating the potential for symbionts to influence STI dynamics. We used laboratory experiments to investigate how a heritable male-killing Spiroplasma bacterial symbiont will affect the epidemiology of a sexually transmitted mite, Coccipolipus hippodamiae in the two-spot ladybird, Adalia bipunctata. We compared mating biology and STI transmission under 1:1 and 4:1 population sex ratios, equivalent to no male-killer presence and 75% male-killer prevalence. We observed that males have the potential to mate with enough females to maintain high female mating rate under strongly female-biased population sex ratios. Further, we observed that females remate more readily when males mate frequently, suggesting female mating rate may in fact be raised in female-biased populations. From the mite's perspective, these effects are countered, at least partly, by reductions in mite transmission from recently mated infected males. We modelled the impact of these processes on STI epidemiology, and concluded that strong sex ratio biases associated with male-killing bacteria are likely to speed the spread of STI infection through both male and female hosts. Thus, the presence of the male-killing bacterium is likely to have indirect impacts on female hosts, mediated through the STI.

Preference for isolated host plants facilitates invasion ofDanaus chrysippus(Linnaeus, 1758) (Lepidoptera: Nymphalidae) by a bacterial male-killerSpiroplasma

Matrilinearly inherited endosymbiotic bacteria are estimated to occur in a third of all terrestrial arthropods. They include male killers, which are typically associated with organisms that lay their eggs in batches. Batch laying favours male-killing bacteria because infected female larvae can cannibalise dead brothers and do not have to compete with them for larval resources. This paper tackles the paradox of high male-killer infection rates in Danaus chrysippus (Linnaeus, 1758), a butterfly that lays its eggs singly rather than in batches, making the selective advantage of male killing for the bacterium hard to understand. The study was conducted in the Nairobi region of Kenya where two substantially allopatric but seasonally migratory semispecies of Danaus chrysippus overlap and hybridise. Within this hybrid zone, but apparently not outside it, D. chrysippus hosts the endosymbiotic male killer, Spiroplasma ixodetis. In Nairobi, this bacterium can infect up to 95% of females. Semispecies D. chrysippus chrysippus and its hybrid with D. chrysippus dorippus (form transiens) are heavily infected, unlike the parental pure D. chrysippus dorippus semispecies, which may be protected by a suppressor gene. We find that: (1) adult females oviposit preferentially on isolated host plants; (2) female larvae on isolated plants survive better than on plants growing in clumps; (3) female larvae survive in greater numbers on plants removed to the laboratory compared to plants left outside; (4) high Spiroplasma infection favours female survival; and (5) high egg/larval density adversely affects adult body size. We conclude that: (1) the butterflies' preference for isolated host plants creates high egg densities and hence enables invasion by Spiroplasma; (2) male killing favours female survival through cannibalism of male eggs and the extra food-plant resources so released; and (3) the butterfly's preference for isolated host plants may provide an enemy-free space that protects eggs and larvae from predators and parasitoids.

Prevalence of male-killer in a sympatric population of two sibling ladybird species, Harmonia yedoensis and Harmonia axyridis (Coleoptera: Coccinellidae)

In predatory ladybirds male embryos in clutches of eggs infected with male-killing bacterial endosymbionts do not hatch and are consumed by female hatchlings. Moreover, it is predicted that infection with male-killing bacteria should be prevalent in a ladybird population if the fitness advantage to female ladybirds due to the resulting reallocation of resources is high. We compared male-killer bacterial prevalence in two sibling species of the ladybird genus Harmonia that use different host ranges. Harmonia yedoensis feeds mostly on the highly elusive pine aphid and its hatchlings can greatly enhance their ability to capture prey and thus their survival by consuming un-hatched eggs in the clutch. In contrast, Harmonia axyridis feeds on a wide range of prey and consumption of un-hatched eggs by the hatchlings does not necessarily increase their foraging success. In the study area where these two species of ladybird occur sympatrically 14 of the 22 females of H. yedoensis (64%) laid clutches of eggs of which approximately only a half hatched and did not produce male offspring, whereas only one of 12 of the females of H. axyridis (8%) exhibited symptoms indicating infection with male-killing bacteria. In the H. yedoensis that exhibited symptoms of being infected with male-killing bacteria, the total allocation of resources to each hatchling, which was provided by laying larger eggs and eating the un-hatched eggs in the clutch, was considerable. These results suggest that the difference in the prevalence of infection with male-killing bacteria in these two species of ladybird may reflect differences in the quality of their prey and their foraging behaviour.

The Japanese ladybirds, Coccinula crotchi and Coccinula sinensis, are infected with very closely related strains of male‐killing Flavobacterium

Male-killing is 1 of 4 known strategies that inherited parasitic endosymbionts have evolved to manipulate their host's reproduction. In early male-killing, infected male offspring are killed early in embryogenesis. Within the Insecta, male-killing bacteria have been found in a wide range of hosts. The Coccinellidae families of beetles, better known as ladybirds, are particularly prone to male-killer invasion. In samples of the coccinellid, Coccinula crotchi, from Japan, a new male-killing bacterium was revealed by phenotypic assay. Molecular genetic analysis revealed the identity to be a tetracycline-sensitive Flavobacterium that causes female-biased offspring sex ratio. Furthermore, that Flavobacterium strain was found to be closely related to the Flavobacterium causing male-killing in the congeneric Japanese coccinellid, Coccinula sinensis, which was collected from the same region. However, we found slightly different Flavobacterium strains infecting C. sinensis from regions with different environmental conditions. This may be an indication of horizontal transmission of male-killing Flavobacterium between these 2 ladybird spices. Finally, environmental conditions may affect the spread of male-killing bacteria among their hosts.

Male Killing Spiroplasma Preferentially Disrupts Neural Development in the Drosophila melanogaster Embryo

Male killing bacteria such as Spiroplasma are widespread pathogens of numerous arthropods including Drosophila melanogaster. These maternally transmitted bacteria can bias host sex ratios toward the female sex in order to ‘selfishly’ enhance bacterial transmission. However, little is known about the specific means by which these pathogens disrupt host development in order to kill males. Here we show that a male-killing Spiroplasma strain severely disrupts nervous tissue development in male but not female D. melanogaster embryos. The neuroblasts, or neuron progenitors, form properly and their daughter cells differentiate into neurons of the ventral nerve chord. However, the neurons fail to pack together properly and they produce highly abnormal axons. In contrast, non-neural tissue, such as mesoderm, and body segmentation appear normal during this time, although the entire male embryo becomes highly abnormal during later stages. Finally, we found that Spiroplasma is altogether absent from the neural tissue but localizes within the gut and the epithelium immediately surrounding the neural tissue, suggesting that the bacterium secretes a toxin that affects neural tissue development across tissue boundaries. Together these findings demonstrate the unique ability of this insect pathogen to preferentially affect development of a specific embryonic tissue to induce male killing.

The direct effects of male killer infection on fitness of ladybird hosts (Coleoptera:Coccinellidae)

Male killing bacteria are common in insects and are thought to persist in host populations primarily by indirect fitness benefits to infected females, whereas direct fitness effects are generally assumed to be neutral or deleterious. Here, we estimated the effect of male killer infection on direct fitness (number of eggs laid, as a measure of fecundity, together with survival) and other life-history traits (development time and body size) in seven ladybird host/male killer combinations. Effects of male killers on fecundity ranged, as expected, from costly to neutral; however, we found evidence of reduced development time and increased survival and body size in infected strains. Greater body size in Spiroplasma-infected Harmonia axyridis corresponded to greater ovariole number and therefore higher potential fecundity. To our knowledge, this is the first report of direct benefits of male killer infection after explicitly controlling for indirect fitness effects. Neutral or deleterious fitness effects of male killer infection should not therefore be automatically assumed.

Two Strains of Male-Killing Wolbachia in a Ladybird, Coccinella undecimpunctata, from a Hot Climate

Ladybirds are a hot-spot for the invasion of male-killing bacteria. These maternally inherited endosymbionts cause the death of male host embryos, to the benefit of female sibling hosts and the bacteria that they contain. Previous studies have shown that high temperatures can eradicate male-killers from ladybirds, leaving the host free from infection. Here we report the discovery of two maternally inherited sex ratio distorters in populations of a coccinellid, Coccinella undecimpunctata, from a hot lowland region of the Middle East. DNA sequence analysis indicates that the male killing is the result of infection by Wolbachia, that the trait is tetracycline sensitive, and that two distinct strains of Wolbachia co-occur within one beetle population. We discuss the implications of these findings for theories of male-killing and suggest avenues for future field-work on this system.

Biased sex ratios and aposematic polymorphism in African butterflies: A hypothesis

In East and Central Africa, the nymphalid butterflies Danaus chrysippus, Acraea encedon and Acraea encedana are involved in a Müllerian mimicry complex. Unusually for aposematic mimetic organisms, the three species show extensive colour pattern polymorphism. Within the same geographic zone, the three species show female-biased sex ratios as a consequence of infection by maternally-inherited, male-killing bacteria. The co-occurrence of biased sex ratios and aposematic polymorphism within these three sympatric, mimetic butterfly species has led to the speculation that invasion by the male-killing bacteria and the subsequent female-biased sex ratios are the underlying reason for the maintenance of colour polymorphism in these species, following its establishment by periods of allopatry and monomorphism and hybridisation. In this paper, we present a novel hypothesis that describes a mechanism by which such causal link might have taken place; in our view, positive frequency-dependent selection favouring the most common colour form in the species is disrupted as a consequence of the recurrent extinction-recolonization cycles undergone at the level of the species populations following the spread of the male-killer. Likewise, extinctions and recolonizations taking place in the other mimics randomly change the direction of selection on each species, potentially leading to multiple selective pressures favouring different colour patterns in the mimicry complex. Thus, selection for monomorphism within each species as well as between the mimetic species will be continuously randomized by the enhanced metapopulation dynamics and the state of polymorphism will be maintained consequently.

The Queen Butterfly, Danaus Chrysippus (L.) (Lepidoptera: Nymphalidae) at Khartoum, Sudan.

Within the region of East and Central Africa, the aposematic butterfly D. chrysippus is characterized by unusual ecology, due to the co-occurrence of polymorphism, mimicry and sex ratio distortion. In this paper, we describe a survey conducted for D. chrysippus at Khartoum, based upon morphological and molecular investigations. The goal of the study was to determine whether Sudan is a part of the presumed hybrid zone of D. chrysippus in Africa, where the butterfly shows the unusual ecology. D. chrysippus was found to display considerable colour pattern polymorphism. Only one mimetic form, the batesian mimic, female Hypolimnas misippus was recorded. The population sex ratio of D. chrysippus did not show significant deviation from the 1:1 ratio. Molecular tests for the male-killing bacteria Spiroplasma yielded negative results. The study concluded that central Sudan belongs to the hybrid zone but lacks its characteristic ecology.

The value of an egg: resource reallocation in ladybirds (Coleoptera: Coccinellidae) infected with male-killing bacteria

Male-killing bacteria are thought to persist in host populations by vertical transmission and conferring direct and/or indirect fitness benefits to their hosts. Here, we test the role of indirect fitness benefits accrued from resource reallocation in species that engage in sibling egg cannibalism. We found that a single-egg meal significantly increased larval survival in 12 ladybird species, but the value of an egg (to survival) differed substantially between species. Next, we tested the impact of three male-killing bacteria on larval survival in one ladybird species, Adalia bipunctata. Spiroplasma reduced larval survival, whereas Wolbachia and Rickettsia had no effect. However, Spiroplasma-infected larvae showed the greatest response to a single-egg meal. The indirect fitness benefit obtained from a single egg is thus so large that even male-killing bacteria with direct fitness costs can persist in host populations. This study supports the hypothesis that fitness compensation via resource reallocation can explain male-killing bacteria persistence.

The Effective Size of Populations Infected With Cytoplasmic Sex-Ratio Distorters

Many arthropod species are infected with maternally inherited endosymbionts that induce a shift in the sex ratio of their hosts by feminizing or killing males (cytoplasmic sex-ratio distorters, or SRDs). These endosymbionts can have profound impacts on evolutionary processes of their hosts. Here, I derive analytical expressions for the coalescent effective size N(e) of populations that are infected with SRDs. Irrespective of the type of SRD, N(e) for mitochondrial genes is given by the number of infected females. For nuclear genes, the effective population size generally decreases with increasing prevalence of the SRD and can be considerably lower than the actual size of the population. For example, with male-killing bacteria that have near perfect maternal transmission, N(e) is reduced by a factor that is given to a good approximation by the proportion of uninfected individuals in the population. The formulae derived here also yield the effective size of populations infected with mutualistic endosymbionts or maternally inherited bacteria that induce cytoplasmic incompatibility, although in these cases, the reduction in N(e) is expected to be less severe than for cytoplasmic SRDs.

Intergenomic Arms Races: Detection of a Nuclear Rescue Gene of Male-Killing in a Ladybird

Many species of arthropod are infected by deleterious inherited micro-organisms. Typically these micro-organisms are inherited maternally. Consequently, some, particularly bacteria of the genus Wolbachia, employ a variety of strategies that favour female over male hosts. These strategies include feminisation, induction of parthenogenesis and male-killing. These strategies result in female biased sex ratios in host populations, which lead to selection for host factors that promote male production. In addition, the intra-genomic conflict produced by the difference in transmission of these cytoplasmic endosymbionts and nuclear factors will impose a pressure favouring nuclear factors that suppress the effects of the symbiont. During investigations of the diversity of male-killing bacteria in ladybirds (Coccinellidae), unexpected patterns of vertical transmission of a newly discovered male-killing taxon were observed in the ladybird Cheilomenes sexmaculata. Initial analysis suggested that the expression of the bacterial male-killing trait varies according to the male(s) a female has mated with. By swapping males between females, a male influence on the expression of the male-killing trait was confirmed. Experiments were then performed to determine the nature of the interaction. These studies showed that a single dominant allele, which rescues male progeny of infected females from the pathological effect of the male-killer, exists in this species. The gene shows typical Mendelian autosomal inheritance and is expressed irrespective of the parent from which it is inherited. Presence of the rescue gene in either parent does not significantly affect the inheritance of the symbiont. We conclude that C. sexmaculata is host to a male-killing γ-proteobacterium. Further, this beetle is polymorphic for a nuclear gene, the dominant allele of which rescues infected males from the pathogenic effects of the male-killing agent. These findings represent the first reported case of a nuclear suppressor of male-killing in a ladybird. They are considered in regard to sex ratio and intra-genomic conflict theories, and models of the evolutionary dynamics and distribution of inherited symbionts.

Spiroplasma infection in Drosophila melanogaster: What is the advantage of killing males?

Male-killing bacteria are maternally inherited agents that cause death of sons of infected females. Their transmission rate is commonly high but imperfect and also sensitive to different environmental factors. Therefore, the proportion of infected females should be reduced in each generation. In order to explain male-killers spread and persistence in host population, a mechanism resulting in the relative increase of infected females must outweigh the losses caused by the imperfect transmission. The resource release hypothesis states that the males’ death results in increased resources available to sibling females which would otherwise be used by their male siblings. Infected females are then expected: to be larger than uninfected females in natural populations; or to have higher viability; or to have shorter development times; or any combination of these outcomes. Here, we tested the resource release hypothesis by measuring body size of infected and uninfected wild-caught Drosophila melanogaster females and carried out other fitness related measures in the laboratory. Wild-caught infected females produced more daughters than uninfected females in their first days in the laboratory. However, although no significant difference in viability was found in a controlled experiment with infected and uninfected flies from a standard laboratory strain, there was a decrease in development time probably mediated by reduced competition. Fitness effects conditioned by the host genetic background are pointed out as a possible explanation for this difference between wild and laboratory flies. Our findings are discussed in the context of the resource advantage hypothesis.