Host selection by the heteronomous hyperparasitoid Encarsia pergandiella: multiple-choice tests using Bemisia argentifolii as primary host

The ovipositional patterns of the heteronomous hyperparasitoid Encarsia pergandiella Howard (Hymenoptera: Aphelinidae) in the presence of its primary host Bemisia argentifolii Bellows & Perring (Hemiptera: Aleyrodidae), and in the presence or absence of conspecific and heterospecific secondary hosts (Encarsia formosa Gahan andEretmocerus mundus Mercet; Hymenoptera: Aphelinidae) were examined to assess host species preferences. Host preferences by heteronomous hyperparasitoids may affect the relative abundance of co-occurring parasitoid species and may influence host population suppression by the parasitoid community. Four combinations of hosts were tested: (1) B. argentifolii, E. mundus, and E. formosa, (2) B. argentifolii, E. formosa, and E. pergandiella, (3) B. argentifolii, E. mundus, and E. pergandiella, and, (4) B. argentifolii, E. mundus, E. formosa, and E. pergandiella. Arrays of hosts (24) were constructed in Petri dishes using leaf disks, each bearing one host. Thirty arrays of each host combination were exposed to single females for 6 h. All hosts were dissected to determine number of eggs per host. Encarsia pergandiella parasitized E. formosa hosts as frequently as E. mundus hosts. However, E. pergandiella parasitized either of these heterospecific hosts more frequently than conspecific hosts in treatments including two secondary host species. When a third parasitoid species was included in host arrays, E. pergandiella parasitized conspecific hosts as frequently as heterospecific hosts. Developmental stage of the hosts did not significantly influence host species selection by E. pergandiella. Our results indicate that host selection and oviposition by heteronomous hyperparasitoids like E. pergandiella, vary with the composition of hosts available for parasitization, and suggest a preference for heterospecific over conspecific secondary hosts.     

Egg Size, Intrinsic Competition, and Lethal Interference in the Parasitoids Encarsia pergandiella and Encarsia formosa

Recent population dynamic theory predicts that disruption of biological control may occur when one parasitoid species' superiority in intrinsic competition is associated with a lower ability to find and exploit hosts (i.e., ability in extrinsic competition). One might expect such a trade-off, for instance, if parasitoids with larger (and fewer) eggs are more likely to prevail in intrinsic competition than species with smaller (and more numerous) eggs. We tested the idea that relative egg size could be used to predict the outcome of intrinsic competition in two closely related endoparasitoids, Encarsia pergandiella Howard and Encarsia formosa Gahan. Contrary to expectation, the parasitoid species with smaller eggs, E. pergandiella, prevailed in intrinsic competition, regardless of the order that hosts were exposed to the two species. In a literature survey, we found four studies of competing pairs of endoparasitoid species for which: (a) egg size estimates were available and (b) one species was consistently superior in intrinsic competition. In three of the four studies, the small-egged species prevailed in intrinsic competition, as we also found. Although E. formosa lost in intrinsic competition, this species negatively affected E. pergandiella's progeny production by host feeding on and killing hosts containing E. pergandiella eggs. E. formosa females also host fed on conspecific-parasitized hosts. As a mechanism of both intra- and interspecific interference competition, host feeding on parasitized hosts contradicts assumptions about the nature of interference competition in existing population dynamics models.     

Cryptic diversity,reproductive isolation and cytoplasmic incompatibility in a classic biological control success story

Molecular genetics and symbiont diagnostics have revolutionized our understanding of insect species diversity, and the transformative effects of bacterial symbionts on host life history. Encarsia inaron is a parasitoid wasp that has been shown to harbour two bacterial endosymbionts, Wolbachia and Cardinium. Known then as E. partenopea, it was introduced to the USA in the late 1980s from populations collected in Italy and Israel for the biological control of an ornamental tree pest, the ash whitefly, Siphoninus phillyreae. We studied natural populations from sites in the USA, the Mediterranean and the Middle East as well as from a Cardinium‐infected laboratory culture established from Italy, with the aims of characterizing these populations genetically, testing reproductive isolation, determining symbiont infection status in their native and introduced range, and determining symbiont role. The results showed that the two Encarsia populations introduced to the USA are genetically distinct, reproductively isolated, have different symbionts and different host–symbiont interactions, and can be considered different biological species. One (‘E. inaron’) is doubly infected by Wolbachia and Cardinium, while only Cardinium is present in the other (‘E. partenopea’). The Cardinium strains in the two species are distinct, although closely related, and crossing tests indicate that the Cardinium infecting ‘E. partenopea’ induces cytoplasmic incompatibility. The frequency of symbiont infection found in the native and introduced range of these wasps was similar, unlike the pattern seen in some other systems. These results also lead to a retelling of a successful biological control story, with several more characters than had been initially described.     

“Darwin's corollary” and cytoplasmic incompatibility induced by Cardinium may contribute to speciation in Encarsia wasps (Hymenoptera: Aphelinidae)

The potential importance of cytoplasmic incompatibility (CI)‐inducing bacterial symbionts in speciation of their arthropod hosts has been debated. Theoretical advances have led to a consensus that a role is plausible when CI is combined with other isolating barriers. However, the insect model systems Nasonia and Drosophila are the only two experimental examples documented. Here, we analyzed the components of reproductive isolation between the parasitoid wasp Encarsia suzannae, which is infected by the CI‐inducing symbiont Cardinium, and its uninfected sibling species Encarsia gennaroi. Laboratory crosses demonstrated that: (1) sexual isolation is incomplete; (2) hybrid offspring production is greatly reduced in the interspecific CI cross; (3) viable hybrids may be produced by curing E. suzannae males of Cardinium with antibiotics; (4) hybrid offspring production in the reciprocal cross is greatly reduced by hybrid inviability due to genetic incompatibilities; (5) hybrid sterility is nearly complete in both directions at the F1 stage. Thus, asymmetrical hybrid incompatibilities and CI act as complementary isolating mechanisms. We propose a new model for contributions of CI symbionts to speciation, with CI reducing gene flow between species in one direction, and in the other, a symbiont sweep resulting in accelerated mtDNA evolution, negative cytonuclear interactions, and hybrid incompatibilities.     

Developmental rates of two congeneric parasitoids,Encarsia formosa and E. pergandiella (Hymenoptera: Aphelinidae), utilizing different egg provisioning strategies

Endoparasitic Hymenoptera vary in the extent to which they provision their eggs and thus in the degree to which they appear to rely on their hosts for resources during embryonic development. In this study, developmental rates were examined in two congeneric parasitoid species, Encarsia formosa and E. pergandiella, that provision their eggs to different degrees. E. formosa eggs are much larger than E. pergandiella eggs. E. formosa eggs hatch significantly earlier than the eggs of E. pergandiella when deposited in 1st or 4th instar nymphs of a common whitefly host, Bemisia tabaci. Both species hatch earlier in 4th instar nymphs, but the delay in hatching in hosts parasitized as 1st instars is much greater in E. pergandiella. While E. formosa develops more rapidly to the 1st larval instar, E. pergandiella emerge as adults significantly earlier, though smaller, than E. formosa adults regardless of the host instar parasitized. These findings show that the extent of provisioning in the eggs of these wasps does not strictly determine their order of progression through different stages of development.     

Secondary host choice by the autoparasitoid Encarsia pergandiella

Autoparasitoids are species of parasitic wasps in the family Aphelinidae which produce females as solitary primary endoparasitoids of homopterans such as whitefly and scale insects (primary hosts), and males as solitary hyperparasitoids. Males generally develop on immature conspecific females or on individuals of other primary parasitoid species (secondary hosts). Encarsia pergandiella is an autoparasitoid that has been introduced to Italy for control of greenhouse whitefly Trialeurodes vaporariorum, in greenhouses and field crops. In this study we examined the secondary host selection behaviour of this species with regard to conspecific females and females of two thelytokous species, E. formosa and E. meritoria. Encarsia formosa has been used successfully for greenhouse whitefly control in Northern Europe, but has not been effective in Southern Italy in winter crops in unheated greenhouses. E. meritoria has recently spread in Italy, and may have potential for biological control of whitefly in the greenhouse environment. In the first experiment, female E. pergandiella were exposed to one of three pair-wise combinations of the three species in petri dish arenas. Parasitism was determined by dissection of the hosts. The number of hosts parasitized by E. pergandiella females did not differ with host species. However, significantly greater numbers of eggs were laid in E. meritoria in both treatments in which it was present; these hosts were more likely to be superparasitized. In a second experiment, observations of females in arenas with equal numbers of all three host species indicated that females encountered and parasitized all host species with approximately equal frequency, although the length of time females spent in the oviposition posture differed with host species.     

Response of adult parasitoids ofBemisia tabaci (Hom.: Aleyrodidae) to leaf residues of selected cotton insecticides

The contact toxicity of eight insecticides to adults of four parasitoids of the sweetpotato whiteflyBemisia tabaci was evaluated in the laboratory. Two common Texas species,Eretmocerus sp. andEncarsia pergandiella Howard, and two exotic species,Eretmocerus mundus Mercet from Spain andEncarsia formosa Gahan from Greece were tested. Insecticides, applied as sprays to greenhouse-grown cotton plants at recommended rates were amitraz (Ovasyn^®), azinphosmethyl (Guthion^®), bifenthrin (Capture^®), buprofezin (Applaud^®), cypermethrin (Ammo^®), methyl parathion and thiodicarb (Larvin^®), with a water control. Parasitoid adults were confined on discs cut from leaves (1) sprayed the same day and (2) sprayed two days previously. Survival in both treatments was measured two and four days following exposure. Significant differences in toxicity were detected among the insecticides. Buprofezin was not toxic to any of the four parasitoids. When caged on leaves sprayed two days previously, only amitraz of the remaining compounds allowed significant general parasitoid survival after two days.E. mundus exhibited the greatest overall tolerance to insecticides, with 40% or more surviving 48 hr after confinement on leaves sprayed with amitraz, thiodicarb and cypermethrin. Survival was generally much reduced after 96 hr. In a separate test, fresh residues of endosulfan (Thiodan^®) were highly toxic at the two rates tested, but two day old residues at the lower rate allowed 76.7% survival ofE. mundus and 35% survival ofE. pergandiella after 48 hrs.     

Poinsettia (Euphorbia pulcherrima Willd Ex Koltz) Cultivar-Mediated Differences in Performance of Five Natural Enemies of Bemisia argentifolii Bellows and Perring, n. sp. (Homoptera: Aleyrodidae)

Laboratory evaluations of five natural enemies of the silverleaf whitefly, Bemisia argentifolii Bellows and Perring, n. sp., were conducted to determine their potential as biological control agents in greenhouse poinsettia ranges. Adult longevity, prey consumption or host feeding and parasitism rates, and parasitoid emergence were measured for one predator, Delphastus pusillus LeConte, and four parasitoids, Encarsia formosa Gahan, Encarsia luteola Howard, Encarsia pergandiella Howard, and Encarsia transvena (Timberlake), as possible indicators of efficacy. Characterization of each parameter was performed on two poinsettia cultivars: the first, ′Annette Hegg Brilliant Diamond,′ has trichome densities on the leaf undersurfaces approximately 15% less than the trichome densities on the leaf undersurfaces of the second cultivar, ′Lilo.′ Adult longevity varied significantly between natural enemies (ranging from an average high of 85.2 days for female D. pusillus feeding on B. argentifolii nymphs to an average low of 2.8 days for the Canada colony of E. formosa), but not between cultivar. Prey consumption and oviposition by D. pusillus varied between prey type (nymphs consumed > eggs consumed) and poinsettia cultivar (′Annette Hegg Brilliant Diamond′ > ′Lilo′). Host feeding, parasitism and total number of B. argentifolii nymphs killed varied significantly among Encarsia spp., but no single wasp performed better than the rest across all three parameters. Host feeding, parasitism, and total number of nymphs killed were greater on ′Annette Hegg Brilliant Diamond′ than on ′Lilo′ and this difference was consistent among the four parasitoid species. Among parasitoid species differences in percentage emergence were consistent between the two poinsettia cultivars with emergence from parasitized nymphs on ′Lilo′ being greater than emergence on ′Annette Hegg Brilliant Diamond.′ Results from these evaluations suggest that the probability of achieving successful augmentative biological central will be greater on poinsettia cultivars with fewer trichomes. In addition, achieving biological control is likely to be difficult with releases of E. transvena, but a greater chance for success may be possible through releases of D. pusillus when whitefly densities are high or through releases of E. formosa (Beltsville colony) or mated E. pergandiella independent of whitefly densities.     

Suitability of stages of femaleEncarsia pergandiella [Hymenoptera: Aphelinidae] for development of conspecific male hyperprasites

Encarsia pergandiella Howard females develop as primary parasitoids of immature whiteflies, whereas ♀♀ develop as secondary parasitoids on ♂♂ of their own species or on other primary whitefly parasitoids. In this experiment, maleE. pergandiella development was examined in the laboratory at 24°C, using different stages of immatureE. pergandiella females [enclosed within the cuticle ofTrialeurodes vaporariorum (Westwood)] as hosts. Unmated adult femaleE. pergandiella were caged individually for 24 h on leaves of plants with either 5-day (early to late 3^rd instar larval), 7-day (late 3^rd instar to prepupal), or 9-day (pupal) hosts. In a control treatment, hosts were allowed to complete development without exposure to unmated adult females. Subsequent emergence of ♂♂ (hosts) and ♀♀ from all treatments was recorded daily. Pupal (9-day)E. pergandiella females were found to be the most suitable hosts for development of ♀♀. Nine- and 7-day hosts were attacked at a similar rate, as indicated by a similarly low proportion of host emergence, but ♀♀ emerged at a significantly higher rate in the 9-day treatment than in the 7-day treatment. Development time of ♀♀ was 15±1 days in the 9-day treatment, and 16±1 days in the 7-day treatment (p<0.005). The proportion of hosts emerging from the 5-day treatment was not significantly different from the control treatment. The data suggest that the colonizing ability ofE. pergandiella in greenhouses is likely to be limited by the delay of 7–9 days between oviposition of female eggs and the suitability of these ♂♂ for oviposition of male eggs. Since development time of the 2 sexes is similar this delay should cause a lack of synchrony in the emergence and mating of F₁ ♂♂ and ♀♀.      

Bemisia tabaci (Homoptera: Aleyrodidae) instar effects on rate of parasitism by Eretmocerus mundus and Encarsia pergandiella (Hymenoptera: Aphelinidae)

Studies were conducted to compare preference among Bemisia tabaci Gennadius, biotype B instars for parasitization by Eretmocerus mundus Mercet and Encarsia pergandiella Howard when provided one instar only, two different instars, and four different instars simultaneously. In the single‐instar no choice treatment, Er. mundus was more successful in parasitizing the younger host instars, while En. pergandiella parasitized a greater proportion of the older instars. Similar results were observed when parasitoids were provided a choice of two instars in six different pair combinations. When all four instars were provided simultaneously, the numbers of first, second, and third instars parasitized by Er. mundus were not significantly different from each other (range 10.3–16.4%), but all were significantly higher than parasitism of fourth instar nymphs (2.1%). The highest percentage parasitization by En. pergandiella was in third instar (17.2%), and the lowest in first instar (2.8%).     

Comparative Genomics Suggests an Independent Origin of Cytoplasmic Incompatibility in Cardinium hertigii

Terrestrial arthropods are commonly infected with maternally inherited bacterial symbionts that cause cytoplasmic incompatibility (CI). In CI, the outcome of crosses between symbiont-infected males and uninfected females is reproductive failure, increasing the relative fitness of infected females and leading to spread of the symbiont in the host population. CI symbionts have profound impacts on host genetic structure and ecology and may lead to speciation and the rapid evolution of sex determination systems. Cardinium hertigii, a member of the Bacteroidetes and symbiont of the parasitic wasp Encarsia pergandiella, is the only known bacterium other than the Alphaproteobacteria Wolbachia to cause CI. Here we report the genome sequence of Cardinium hertigii cEper1. Comparison with the genomes of CI–inducing Wolbachia pipientis strains wMel, wRi, and wPip provides a unique opportunity to pinpoint shared proteins mediating host cell interaction, including some candidate proteins for CI that have not previously been investigated. The genome of Cardinium lacks all major biosynthetic pathways but harbors a complete biotin biosynthesis pathway, suggesting a potential role for Cardinium in host nutrition. Cardinium lacks known protein secretion systems but encodes a putative phage-derived secretion system distantly related to the antifeeding prophage of the entomopathogen Serratia entomophila. Lastly, while Cardinium and Wolbachia genomes show only a functional overlap of proteins, they show no evidence of laterally transferred elements that would suggest common ancestry of CI in both lineages. Instead, comparative genomics suggests an independent evolution of CI in Cardinium and Wolbachia and provides a novel context for understanding the mechanistic basis of CI.     

Induction of males in thelytokous populations of Encarsia meritoria and Encarsia protransvena: A systematic tool

Males from thelytokous females of Italian uniparentalpopulations of Encarsia meritoria and Encarsiaprotransvena were obtained by feeding wasps with tetracycline orrearing parasitoids at high temperatures respectively. Two thermictreatments were tested on E. protransvena: 31°C constanttemperature versus an alternation of 12 hours at 33°C and 23°Con a daily basis. Females of E. meritoria fed with tetracyclineproduced 80.87% of male progeny. Tetracycline did not induce maleoffspring in E. protransvena. For this species hightemperatures were required even though only a few males were obtained(0.31–5.43%). These results confirm the infection ofparthenogenesis-inducing micro-organisms in E. meritoria, whilethis is doubtful in E. protransvena. Males derived fromthelytokous populations of the two species were described. Males of theuniparental population of E. meritoria were similar to those ofbiparental ones, suggesting that the two populations are conspecific.Males of E. protransvena are slightly different from thosedescribed previously, supporting the hypothesis that E.protransvena is a complex of cryptic species. Our results suggestthe possibility of using ``induced males' by either antibiotics or hightemperatures as a taxonomic tool in the genus Encarsia toseparate similar (if not undistinguishable) uniparentalspecies.     

In situ observation of the <Emphasis Type="Italic">Cardinium</Emphasis> symbionts of <Emphasis Type="Italic">Brevipalpus</Emphasis> (Acari: Tenuipalpidae) by electron microscopy

Brevipalpus (Acari: Tenuipalpidae) mites are important pests on a variety of host plant species. The mites damage their hosts directly by feeding and some species also serve as vectors of plant viruses. Among more than 200 described Brevipalpus species, three are recognized as vectors of plant viruses: B. phoenicis, B. californicus and B. obovatus. These species occur worldwide in subtropical and tropical regions. Brevipalpus mites reproduce mostly by thelytokous parthenogenesis and this condition was attributed to a bacterial endosymbiont, recently characterized as a member of the genus Cardinium. The same symbiont infects many other arthropods and is capable of manipulating their host reproduction in various ways. Generally the presence of Cardinium is determined by molecular, PCR based, techniques. In the current work we present visual evidence for the presence of these bacteria by transmission electron microscopy as a complement of previous detection by PCR. Cardinium is easily identified by the presence of a unique array of microtubule-like structures (ML) in the cell. Symbionts have been observed in several organs and eggs from different populations of all three Brevipalpus species known as vector of plant viruses. Cardinium cells were always immersed directly within the cytoplasm of infected cells. Bacteria were observed in all females of all instars, but were absent from all males examined. Females from some Brevipalpus populations were observed to be uninfected by Cardinium. This observation confirmed previous PCR-based results that these populations were aposymbiotic. The observed distribution of the bacteria suggests that these bacteria could have other functions in the mite biology beside feminization.     

A new species ofEncarsia [Hym.: Aphelinidae] parasitizingBemisia tabaci[Hom.: Aleyrodidae]

Encarsia deserti n.sp., a parasite ofBemisia tabaci Gennadius is described. It resemblesE. formosa Gahan, but has a light brown head and thorax, is smaller, and is biparental. This species was found in Southern California and Arizona parasitizingB. tabaci and was introduced into Israel for the control of this pest.     

Endosymbiont costs and benefits in a parasitoid infected with both Wolbachia and Cardinium

Theory suggests that maternally inherited endosymbionts can promote their spread and persistence in host populations by enhancing the production of daughters by infected hosts, either by improving overall host fitness, or through reproductive manipulation. In the doubly infected parasitoid wasp Encarsia inaron, Wolbachia manipulates host reproduction through cytoplasmic incompatibility (CI), but Cardinium does not. We investigated the fitness costs and/or benefits of infection by each bacterium in differentially cured E. inaron as a potential explanation for persistence of Cardinium in this population. We introgressed lines infected with Wolbachia, Cardinium or both with the cured line to create a similar genetic background, and evaluated several parasitoid fitness parameters. We found that symbiont infection resulted in both fitness costs and benefits for E. inaron. The cost was lower initial egg load for all infected wasps. The benefit was increased survivorship, which in turn increased male production for wasps infected with only Cardinium. Female production was unaffected by symbiont infection; we therefore have not yet identified a causal fitness effect that can explain the persistence of Cardinium in the population. Interestingly, the Cardinium survivorship benefit was not evident when Wolbachia was also present in the host, and the reproduction of doubly infected individuals did not differ significantly from uninfected wasps. Therefore, the results of our study show that even when multiple infections seem to have no effect on a host, there may be a complex interaction of costs and benefits among symbionts.     

Parasrnzatlon of bemisia argentifolii (Hom.: Aleyrodidae) by encarsia pergandiella (Hym.: Apheunidae)

The biology of the arrhenotokous autoparasitoid,Encarsia pergandiella Howard, was studied in the laboratory on the silverleaf whitefly,Bemisia argentifolii Bellows & Perring. Egg to adult development of parasitoid females averaged ca. 14 days at about 25.3+0.2?C regardless of whether the whitefly host was reared on tomato, eggplant or squash. While all instars ofB. argentifolii were accepted for primary parasitization, a greater percentage of third and fourth instars were parasitized. Mortality of whitefly nymphs in the absence of parasitization did not differ among instars and averaged about 35%. Second instar to pupal parasitoid females were accepted for secondary parasitization although a greater percent of pupal females were parasitized. About 40% of immatureE. pergandiella females more than 4 days old died in the absence of secondary parasitization when exposed to adultE. pergandiella females.     

Cytoplasmic incompatibility and multiple symbiont infection in the ash whitefly parasitoid, Encarsia inaron

Terrestrial arthropods are commonly infected with maternally inherited symbionts that cause reproductive incompatibilities between hosts with differing infection status. Such symbionts can have major effects on the efficacy of a biological control program if releases are comprised of mixtures of differentially infected individuals. In this study, the ash whitefly parasitoid, Encarsia inaron (Hymenoptera: Aphelinidae) from Arizona was surveyed for the presence of heritable bacterial symbionts; experiments were also performed to test for two phenotypes known to be caused by Encarsia symbionts—cytoplasmic incompatibility and changes in oviposition behavior and host use. E. inaron has successfully reduced ash whitefly to non-pest status in all three locations it has been released (California, Arizona, and North Carolina) and is also notable as one of the only Encarsia species that is not autoparasitic, with both male and female wasps developing as primary parasitoids of whiteflies. We show that E. inaron is infected with both Wolbachia and Cardinium. While there was no effect of the symbionts on oviposition behavior or host use, crosses between doubly infected male wasps and uninfected females resulted in a severe reduction in the number of female offspring; male offspring production was unaffected. This study thus serves as a further warning that ascribing a phenotype to a symbiont with confidence depends on eliminating the possibility of a mixed infection, and establishes E. inaron as a useful model for dissecting Wolbachia–Cardinium interactions.     

Description and identification of two new diseases of guariroba palm (Syagrus oleraceae) in Brazil

Guariroba is a palm species native to central Brazil. Seedlings of guariroba with leaf spots of unknown aetiology were found in Patos de Minas, Minas Gerais, Brazil. The leaf spots were manifest as two different symptom types: the first lesion type consisted of necrotic spots with a rounded to elongate shape, with a light colour and dark edges; the second lesion type had a rounded shape, was dark brown in colour with a light brown edge. The objective of this study was to elucidate the aetiology of both diseases. The likely causal agents were isolated and Koch's postulate fulfilled. Subsequently, the ITS region of rDNA from both micro‐organisms were amplified and sequenced. According to the morphological characteristics and molecular analyses, the fungal species were identified as Pestalotiopsis adusta (causing necrotic spots with a rounded to elongate shape, with a light colour and dark edges), and Alternaria tenuissima (causing lesions with a rounded shape, dark brown in colour with a light brown edge). Identification of the causal organisms of these two diseases will help guide management approaches that might be tested to reduce impact of the disease on Guariroba, including the use of fungicides and cultural approaches.     

Biology and distribution of microbe-associated thelytokous populations of aphid parasitoids (Hym., Braconidae, Aphidiinae)

Abstract: Some phenomena of the biology and ecology of the microbe ( Wolbachia )-associated thelytokous populations of aphid parasitoids are presented for the first time. Thelytokous virgin females were found to refuse the mating attempts by males of biparental populations, although males occurred quite rarely among the offspring. The host range was found to be about identical in conspecific biparental and thelytokous populations, whereas the association of solely biparental populations with a certain host group might be considered as a marker of the presence of sibling species in the area. The relative frequency of conspecific biparental and thelytokous populations varied, depending upon the area and host species. The microbe-induced thelytokous populations were confirmed in three species of the genus Lysiphlebus Foerst. ( Lysiphlebus cardui Marsh., Lysiphlebus confusus Tremblay and Eady, Lysiphlebus fabarum Marsh.) and probably some other, as yet undescribed, species only in the West Palearctic subregion, whereas the other known congeneric species in the subregion as well as in the Nearctic and the Neotropical regions were determined as strictly biparental species. Microbe-induced thelytokous populations did not change due to host species alternation and/or distribution area (purposeful introductions as biocontrol agents).     

Epidemiology of asexuality induced by the endosymbiotic Wolbachia across phytophagous wasp species: host plant specialization matters

Among eukaryotes, sexual reproduction is by far the most predominant mode of reproduction. However, some systems maintaining sexuality appear particularly labile and raise intriguing questions on the evolutionary routes to asexuality. Thelytokous parthenogenesis is a form of spontaneous loss of sexuality leading to strong distortion of sex ratio towards females and resulting from mutation, hybridization or infection by bacterial endosymbionts. We investigated whether ecological specialization is a likely mechanism of spread of thelytoky within insect communities. Focusing on the highly specialized genus Megastigmus (Hymenoptera: Torymidae), we first performed a large literature survey to examine the distribution of thelytoky in these wasps across their respective obligate host plant families. Second, we tested for thelytoky caused by endosymbionts by screening in 15 arrhenotokous and 10 thelytokous species for Wolbachia, Cardinium, Arsenophonus and Rickettsia endosymbionts and by performing antibiotic treatments. Finally, we performed phylogenetic reconstructions using multilocus sequence typing (MLST) to examine the evolution of endosymbiont‐mediated thelytoky in Megastigmus and its possible connections to host plant specialization. We demonstrate that thelytoky evolved from ancestral arrhenotoky through the horizontal transmission and the fixation of the parthenogenesis‐inducing Wolbachia. We find that ecological specialization in Wolbachia's hosts was probably a critical driving force for Wolbachia infection and spread of thelytoky, but also a constraint. Our work further reinforces the hypothesis that community structure of insects is a major driver of the epidemiology of endosymbionts and that competitive interactions among closely related species may facilitate their horizontal transmission.