The influence of temperature and host availability on the host exploitation strategies of sexual and asexual parasitic wasps of the same species

In the hymenopteran parasitoid Venturia canescens, asexual (obligate thelytoky not induced by Wolbachia bacteria) and sexual (arrhenotokous) wasps coexist in field conditions despite the demographic cost incurred due to the production of males by sexual females. Arrhenotoky predominates in field conditions, whereas populations in indoor conditions (mills, granaries) are exclusively thelytokous. These differences in the relative abundance of the two modes of reproduction between environments suggest that the individuals of each reproductive mode may have developed strategies adapted to the conditions prevailing in each kind of habitat. The two environments contrast in temperature variability and in the spatial heterogeneity of host availability. In this study, we considered the combined effect of temperature and host availability on host patch exploitation by thelytokous and arrhenotokous V. canescens. As expected, arrhenotokous females were more sensitive to temperature changes. If the temperature decreased before foraging, they remained longer and exploited patches more thoroughly. This is consistent with the expected behaviour of parasitoids in response to signs of unfavourable conditions that entail increasing risk of time limitation or a reduced probability of attaining further patches. Both arrhenotokous and thelytokous females increased patch exploitation with host availability. However, unexpectedly, we found no difference in the way the two types of wasp responded to differences in host availability. Differences in the strategies adopted under different environmental conditions may indicate divergence of niche-specific life history traits between the two modes of reproduction. Niche displacement may partly account for the coexistence of these two modes of reproduction at a geographical scale.     

Differential energy allocation as an adaptation to different habitats in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Environmental pressures are expected to favour organisms that optimally allocate metabolic resources to reproduction and survival. We studied the resource allocation strategies and the associated tradeoffs in the parasitoid wasp Venturia canescens, and their adaptation to the characteristics of the environment. In this species, individuals of two reproductive modes coexist in the same geographical locations, but they mainly occur in distinct habitats. Thelytokous (asexual) wasps are mostly found in anthropogenic habitats, where hosts tend to aggregate and food is absent. Arrhenotokous (sexual) wasps are exclusively found in natural habitats, where hosts are scattered and food is present. We analysed (1) the quantity of energy stored during ontogeny, (2) the tradeoff between reproduction and survival, by measuring egg load and longevity and (3) the host patch exploitation behaviour of the wasps at emergence. Arrhenotokous wasps emerged with more metabolic resources than thelytokous ones, especially glycogen, a nutrient that could be used for flying in search of hosts and/or food. Thelytokous wasps allocated more energy than arrhenotokous wasps to egg production: this would allow them to parasitize more hosts. The tradeoff between egg production and longevity was not revealed within reproductive modes, but when comparing them. At emergence, arrhenotokous wasps tended to exploit host patches less thoroughly than thelytokous wasps, suggesting that by leaving the host patch, they search for food. The results clearly showed adaptations to the characteristics of habitats preferentially inhabited by the two reproductive modes, and suggested a mechanism that facilitates their coexistence in natural conditions.     

Detection of Gene Flow from Sexual to Asexual Lineages in Thrips tabaci (Thysanoptera: Thripidae)

Populations of Thrips tabaci are known to have two sympatric but genetically isolated reproductive modes, arrhenotoky (sexual reproduction) and thelytoky (asexual reproduction). Herein, we report behavioral, ecological and genetic studies to determine whether there is gene flow between arrhenotokous and thelytokous T. tabaci. We did not detect significant preference by arrhenotokous males to mate with females of a particular reproductive mode, nor did we detect significant behavioral differences between arrhenotokous males mated with arrhenotokous or thelytokous females in their pre-copulation, copulation duration and mating frequency. Productive gene transfer resulting from the mating between the two modes was experimentally confirmed. Gene transfer from arrhenotokous T. tabaci to thelytokous T. tabaci was further validated by confirmation of the passage of the arrhenotokous male-originated nuclear gene (histone H3 gene) allele to the F₂ generation. These behavioral, ecological and genetic studies confirmed gene transfer from the sexual arrhenotokous mode to the asexual thelytokous mode of T. tabaci in the laboratory. These results demonstrate that asexual T. tabaci populations may acquire genetic variability from sexual populations, which could offset the long-term disadvantage of asexual reproduction.     

Gene flow between arrhenotokous and thelytokous populations of Venturia canescens (Hymenoptera)

In the solitary parasitoid wasp Venturia canescens both arrhenotokously (sexual) and thelytokously (parthenogenetical) reproducing individuals occur sympatrically. We found in the laboratory that thelytokous wasps are able to mate, receive and use sperm of arrhenotokous males. Using nuclear (amplified fragment length polymorphism, virus-like protein) and mitochondrial (restriction fragment length polymorphism) markers, we show the occurrence of gene flow from the arrhenotokous to the thelytokous mode in the field. Our results reinforce the paradox of sex in this species.     

Biological Control Potential of Wolbachia -infected Versus Uninfected Wasps: Laboratory and Greenhouse Evaluation of Trichogramma cordubensis and T. deion Strains

The effects of thelytoky-inducing Wolbachia ( &#102 -proteobacteria) on Trichogramma cordubensis and T. deion (Hymenoptera, Trichogrammatidae) were studied in laboratory and greenhouse conditions. One infected (thelytokous, all female) line of each wasp species was compared with its conspecific uninfected (arrhenotokous, sexual) counterpart for several fecundity and dispersal traits. Arrhenotokous lines had a higher fecundity than their thelytokous counterparts, which suggests that Wolbachia negatively affect the fecundity of Trichogramma females. The arrhenotokous females dispersed more in the laboratory than their thelytokous counterparts. In the greenhouse, the opposite effect or no difference between lines was found, indicating that the laboratory set-up used to measure dispersal is not useful to predict relative dispersal of the females in the greenhouse. Calculations show that by releasing 100 adult wasps of both lines, thus including arrhenotokous males in the sexual line, more eggs are parasitized by the thelytokous wasps. Therefore, in spite of their lower individual female fecundity, thelytokous lines have a better potential for biological control than their arrhenotokous counterparts.     

Mitochondrial DNA diversity and geographical distribution of sexual and asexual strains of the braconid parasitoid Meteorus pulchricornis

Asexuality is an important tool with regard to the use of parasitoid wasps as biocontrol agents. Asexual (apomictic thelytokous) strains of Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae), a polyphagous endoparasitoid of lepidopteran larvae, are sympatric with sexual (arrhenotokous) strains in Japan. The results of phylogenetic analyses of mitochondrial cytochrome c oxidase subunit I (COI) sequences indicated two major haplotype groups on the Japanese islands. The northern group I predominantly contained sexual strains, whereas the southern group II contained both sexual and asexual strains. Most asexual strains were likely derived within group II. An asexual strain recently established in New Zealand has the identical haplotype to a strain in Japan and was proven to have originated from East Asia. Three hypotheses on the evolution of asexuality are discussed for this parasitoid wasp: recessive gene, hybridization, and cytoplasmic element.     

Reduced sexual functionality of PI‐Wolbachia‐infected females of Tetrastichus coeruleus

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts by, for example, inducing parthenogenesis. In most cases of Wolbachia‐induced parthenogenesis, the infection is fixed and the entire host population consists of females. In the absence of males and sexual reproduction, genes involved in sexual reproduction are not actively maintained by selection. Accumulation of neutral mutations or selection against maintenance of sexual traits may lead to their loss or deterioration. In addition, females may lose the ability to reproduce sexually due to ‘functional virginity mutations’ that may spread concomitantly with the Wolbachia infection through a population. The parasitoid wasp Tetrastichus coeruleus (Nees) (Hymenoptera: Eulophidae) forms an ideal model to study the decay of sexual functionality, because it has both Wolbachia‐infected, parthenogenetic populations and uninfected, sexual populations. We compared several components of sexual functionality of arrhenotokous (sexual) and thelytokous (parthenogenetic) T. coeruleus females. First, we tested whether arrhenotokous and thelytokous females were equally attractive and receptive to males. Second, we examined whether mating is costly to females by measuring the life span of mated and virgin females. Last, we studied the morphology of the spermathecae of arrhenotokous and thelytokous females. Mated females had shorter life spans than virgin females, showing that mating carried a fitness cost. Two sexual traits of thelytokous females have degraded compared to arrhenotokous females. Arrhenotokous and thelytokous females were equally attractive to males, but thelytokous females were unreceptive to males. Furthermore, there was a clear difference in spermathecal morphology between arrhenotokous and thelytokous females. Our data do not allow distinction between the various potential causes of such degradation. Although the longevity cost of mating may indicate selection against the maintenance of costly sexual traits, accumulation of neutral mutations, functional virginity mutations, manipulation by Wolbachia, and/or the genetic distance between the two populations may all have contributed to the decay of sexual traits in thelytokous females.     

Are individuals from thelytokous and arrhenotokous populations equally adept as biocontrol agents? Orientation and host searching behavior of a fruit fly parasitoid

Hymenopteran parasitoids generally reproduce by arrhenotoky, in which males develop from unfertilized eggs and females from fertilized eggs. A minority reproduce by thelytoky, in which all-female broods are derived from unfertilized eggs. Thelytokous populations are potentially of interest for augmentative biological control programs since the exclusive production of females could significantly lower the costs of mass rearing. Behavioral traits are a major component of parasitoid efficacy. Here, we examined orientation and host searching behavior in thelytokous and arrhenotokous populations of the fruit fly parasitoid Odontosema anastrephae Borgmeier (Hymenoptera: Figitidae). Orientation behavior to various odorant sources was studied in a two-choice olfactometer. No major differences were found between thelytokous and arrhenotokous wasps for this behavior. However, when host-searching behaviors were analyzed, some differences were found. Thelytokous females arrived sooner, foraged longer, and remained longer on non-infested guavas than arrhenotokous females. Individuals of both forms exhibited similar stereotyped behavioral sequences vis-à-vis guava treatments, with only slight deviations detected. Our results suggest that individuals from selected thelytokous and arrhenotokous O. anastrephae populations have similar abilities to search for tephritid larvae, supporting the use of thelytokous strains for augmentative releases.     

Could different environmental conditions select for different flight characteristics in sexual and asexual parasitoid Venturia canescens?

In the parasitoid Venturia canescens Gravenhorst (Hymenoptera: Ichneumonidae), asexual and sexual wasps coexist in the field in the Mediterranean basin, but only the asexual strain is present indoors. The sexual strain dominates outdoors despite the demographic costs associated with the production of males and mate location. The present study tests whether females of the sexual and asexual strains of V. canescens differ in flight characteristics in line with the differences of their preferred habitats and enquires whether these differences might contribute to the persistence of sexually reproducing individuals in the vicinity of asexual counterparts. The results show that sexual female wasps are smaller than their asexual counterparts. The size of wasps has a strong influence on flight parameters, with larger animals generally being better fliers. In wasps of approximately the same size, sexual wasps fly faster than their asexual counterparts under experimental laboratory conditions, in terms of both the average speed over the observation period as well as the longest single flight. Sexual wasps also perform fewer flights to cover the same distance. Sexual wasps have higher wing loading than asexual ones of the same size, which could have contributed to the observed differences in speed between individuals of both reproductive modes. There are no significant differences between the two reproductive modes in the parameters related to the distance traversed or the time spent in flight. This study shows clear differences in the flight behaviour of sexual and asexual V. canescens. Together with previous results, this finding suggests differential adaptations to their preferred habitats. These differences might ease the competition between modes of reproduction through niche and habitat differentiation and might help to explain their coexistence on a geographical scale.     

The dynamics of energy allocation in adult arrhenotokous and thelytokous Venturia canescens

The ability to adjust resource allocation to the quality of the environment has broad implications for animal reproductive success. Organisms with complex life cycles that may experience various selection pressures during their lifetime are expected to evolve mechanisms to modulate the resource allocation strategies adopted during ontogeny to the conditions encountered by the adult. In the parasitoid Venturia canescens Gravenhorst (Hymenoptera: Ichneumonidae), thelytoky appears to have been selected for in anthropogenic habitats, where hosts are relatively numerous and food is absent, and arrhenotoky in natural habitats where hosts are more scarce and food is present. A previous study postulated that during their juvenile stage, females of both reproductive modes adopt strategies of energy allocation in accordance with these conditions, possibly providing a direct short‐term advantage to arrhenotokous forms, which partially co‐occur with thelytokous forms under natural conditions. To test this assumption, we provided daily adult thelytokous and arrhenotokous females with a small number of hosts together with food. To compare their lifetime resource allocation strategies, we recorded wasp longevities, egg loads, and carbohydrate reserves in wasps of different ages. Our analysis indicates that thelytokous females are able, to a certain extent, to cope with these conditions, because they reached the same longevity as arrhenotokous females. Nevertheless, thelytokous females suffered from a higher degree of time limitation compared with arrhenotokous ones, and arrhenotokous wasps appeared to maintain their energetic advantage over the adult stage. These results provide new insights, and point to the consideration of other activities, such as flight performance and/or ability to reach food and hosts, in the understanding of the role of resource allocation strategies in the maintenance of sex in this species.     

Mating system and population structure in the desert ant Cataglyphis livida

We investigated population genetic structure, mating system, worker reproduction and thelytokous parthenogenesis in the desert ant Cataglyphis livida. Pedigree analyses at polymorphic microsatellite loci show that colonies are headed by a single queen, and that queens are mated with two to eight males. No inbreeding was found in the population sampled. Colonies are genetically differentiated and exhibit no isolation-by-distance pattern, consistent with independent foundation of new colonies. Workers do reproduce and lay haploid (arrhenotokous) eggs in queenless colonies; conversely, we found no evidence of worker reproduction in queenright nests. In contrast with C. cursor, where new queens are produced by thelytokous parthenogenesis, female sexuals and workers of C. livida arise from classical sexual reproduction. We discuss the parallels and contrasts between the mating system and population structure in C. livida and the other Cataglyphis species studied so far.     

The cytological basis for reproductive variability in the Anoetidae (Sarcoptiformes: Acari)

Meiosis is described in a thelytokous strain of the anoetid mite Histiostoma feroniarum (Dufour) and in both sexes of the arrhenotokous strain of this species. Oogenesis in the thelytokous strain is accomplished by ameiotic mitosis with only one pseudo-maturation division. During this division one or more chromosomes may move to the poles precociously and while in this position can be mistaken for centrioles. Fourteen chromosomes are found at metaphase of the pseudo-maturation division and in cleaving eggs of this strain. In the arrhenotokous strain, male meiosis consists of a single mitotic division. Oogenesis is regular and 7 bivalents are observed at the first maturation division. Metaphases of the first cleavage division in fertilized eggs show 14 chromosomes and 7 chromosomes in unfertilized eggs.It is postulated that the thelytokous strain has arisen from the arrhenotokous strain. This assumption is in agreement with that suggested for several insect species previously reported. The evolution in the Acari and the variability in the modes of reproduction in this suborder are discussed in light of the findings in this paper on the Anoetidae.     

MOLECULAR MARKERS INDICATE RARE SEX IN A PREDOMINANTLY ASEXUAL PARASITOID WASP

The parasitoid wasp genus Lysiphlebus (Hymenoptera: Braconidae: Aphidiinae) contains a taxonomically poorly resolved group of both sexual (arrhenotokous) species and asexual (thelytokous) clones. Maximum-parsimony and maximum-likelihood analyses of mitochondrial DNA sequence data from specimens collected across Western Europe showed that asexuality, which does not appear to be caused by the bacterium Wolbachia, is concentrated in two geographically widespread lineages, the older of which diverged from the closest extant sexual taxa approximately 0.5 million years ago. However, the DNA sequences of a nuclear intron (elongation factor—1α) showed no congruence with this pattern, and a much higher frequency of heterozygotes with very high allelic diversity was observed among the asexual females compared to that among females from the sexual species. This pattern is consistent with maternally inherited asexuality coupled with a history of rare sex with members of several closely related sexual populations or species. Our observations reinforce recent arguments that rare sex may be more important for the persistence of otherwise asexual lineages than hitherto appreciated.     

Odor-mediated patch choice in the parasitoid Venturia canescens: temporal decision dynamics

Parasitoids foraging for hosts in a heterogeneous environment would greatly benefit if they could decide already from a distance in which areas search for resources would be most profitable and to avoid areas of low fitness returns. Interestingly, the temporal dynamics of the decision process in parasitoid patch choice have rarely been investigated. In a Y-tube olfactometer, we tested whether thelytokous and arrhenotokous females of the parasitoid wasp Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae) respond to differences in cues indicating the quality of a host-containing patch and choose more profitable patches. Special attention was given to the time it took females to make their choices (patch choice time) when differences in patch quality were either qualitative (absence vs. presence of hosts and kairomone) or quantitative (various concentrations of hosts and kairomone, and presence of competitors). We found that both thelytokous and arrhenotokous wasps only chose the higher-quality patch based on odor cues when the difference was qualitative. When patches differed only with respect to the number of hosts, or the presence or absence of competing female parasitoids, no significant preference could be found in females of either strain of the parasitoid. In contrast, both the time until females reached the junction of the Y-tube olfactometer (response time) and the time until females decided for either patch (decision time) varied with parasitoid strain and odor treatment. Thelytokous wasps were faster than arrhenotokous wasps in their response time and in their decision time. However, females of both strains responded faster with increasing number of total hosts releasing kairomone. Yet, decision time for patches did not significantly vary as a function of patch quality offered to Venturia wasps.     

No evidence of parthenogenesis‐inducing bacteria involved in Thripoctenus javae thelytoky: an unusual finding in Chalcidoidea

All Hymenoptera have a haplodiploid mode of sex determination. Although most species reproduce by arrhenotokous parthenogenesis, there are many thelytokous species, in which unfertilized eggs develop into diploid females. Thelytoky can be genetic or due to microbial infection. In the large Chalcidoidea superfamily, thelytokous parthenogenesis is almost always associated with infection of endosymbionts of the genera Wolbachia, Cardinium, and Rickettsia. Thripoctenus javae (Girault) (Hymenoptera: Eulophidae) is a larval parasitoid of the greenhouse thrips Heliothrips haemorrhoidalis (Bouché) (Thysanoptera: Thripidae), an important worldwide pest. Both the host and its parasitoid reproduce by thelytokous parthenogenesis. The main goal of this study was to test whether endosymbiotic bacteria, either those known to induce thelytokous parthenogenesis or other sex‐manipulators, are responsible for thelytoky of two geographically distinct populations of T. javae. We used sequencing of ribosomal ITS2 and 28S‐D2 and mitochondrial COI genes to molecularly characterize the two populations, antibiotic and heat treatments, and FISH of ovaries, for thelytoky studies. It was impossible to revert thelytokous individuals back to sexual reproduction and no evidence of bacterial infection was found in parthenogenetic T. javae females. This makes T. javae the second chalcidoid in which thelytokous reproduction appears not to be associated with the presence of bacterial endosymbionts.     

Population genetics of Wolbachia‐infected,parthenogenetic and uninfected,sexual populations of Tetrastichus coeruleus (Hymenoptera: Eulophidae)

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts. These manipulations are expected to have consequences on the population genetics of the host, such as heterozygosity levels, genetic diversity and gene flow. The parasitoid wasp Tetrastichus coeruleus has populations that are infected with parthenogenesis‐inducing Wolbachia and populations that are not infected. We studied the population genetics of T. coeruleus between and within Wolbachia‐infected and uninfected populations, using nuclear microsatellites and mitochondrial DNA. We expected reduced genetic diversity in both DNA types in infected populations. However, migration and gene flow could introduce new DNA variants into populations. We therefore paid special attention to individuals with unexpected (genetic) characteristics. Based on nuclear and mitochondrial DNA, two genetic clusters were evident: a thelytokous cluster containing all Wolbachia‐infected, parthenogenetic populations and an arrhenotokous cluster containing all uninfected, sexual populations. Nuclear and mitochondrial DNA did not exhibit concordant patterns of variation, although there was reduced genetic diversity in infected populations for both DNA types. Within the thelytokous cluster, there was nuclear DNA variation, but no mitochondrial DNA variation. This nuclear DNA variation may be explained by occasional sex between infected females and males, by horizontal transmission of Wolbachia, and/or by novel mutations. Several females from thelytokous populations were uninfected and/or heterozygous for microsatellite loci. These unexpected characteristics may be explained by migration, by inefficient transmission of Wolbachia, by horizontal transmission of Wolbachia, and/or by novel mutations. However, migration has not prevented the build‐up of considerable genetic differentiation between thelytokous and arrhenotokous populations.     

Evidence that mass trapping suppresses pink bollworm populations in cotton fields

Both thelytokous and arrhenotokous Trichogramma minutum were collected from eggs of the spruce budmoth, Zeiraphera canadensis in New Brunswick, Canada and their phenotypic traits compared for use in biological control. The lower threshold temperature for development of thelytokous and arrhenotokous parasitoids was 9.7 and 10.1 °C, respectively; the former required significantly higher degree-days for development from egg to adult (–x±SE=165.1±5.8 °D) than the latter (128.1±4.9 °D). Thelytokous and arrhenotokous parasitoids had similar forewing length (0.49±0.01 vs. 0.49±0.01 mm) and adult lifespan (13.3±0.7 vs. 14.0±1.1 days), but significantly different fecundity and sex ratios. Thelytokous females produced fewer offspring (89.5±6.6 vs. 173.9±6.4) and fewer female progeny (77.2±5.4 vs. 109.8±3.3) despite an overall higher proportion of females (91.6±1.1% vs. 65.4±2.8%) than their arrhenotokous counterparts. Ovarian dissections showed that the number of eggs increased with parasitoid age in arrhenotokous parasitoids but remained steady in thelytokous parasitoids. The variation in ovarian development of the two forms was the major factor contributing to the differences in fecundity. Thelytokous parasitoids were more host-specific than arrhenotokous ones; when offered eight host species, the former rejected three whereas the latter rejected only one. Thelytokous parasitoids survived better than arrhenotokous ones when stored from 30 to 150 days at 4 °C. Thelytokous females were slower at initiating flight after emergence than arrhenotokous females but maintained flight activity longer (6 h). These results indicate that thelytokous T. minutum are different from their arrhenotokous counterparts physiologically, biologically and ecologically and that they may play different roles in the field. The potential for using thelytokous parasitoids in biological control programs is discussed.     

A single locus determines thelytokous parthenogenesis of laying honeybee workers (Apis mellifera capensis)

The evolution and maintenance of parthenogenetic species are a puzzling issue in evolutionary biology. Although the genetic mechanisms that act to restore diploidy are well studied, the underlying genes that cause the switch from sexual reproduction to parthenogenesis have not been analysed. There are several species that are polymorphic for sexual and parthenogenetic reproduction, which may have a genetic basis. We use the South African honeybee subspecies Apis mellifera capensis to analyse the genetic control of thelytoky (asexual production of female workers). Due to the caste system of honeybees, it is possible to establish classical backcrosses using sexually reproducing queens and drones of both arrhenotokous and thelytokous subspecies, and to score the frequency of parthenogenesis in the resulting workers. We found Mendelian segregation for thelytoky of egg-laying workers, which appears to be controlled by a single major gene (th). The segregation pattern indicates a recessive allele causing thelytoky. We found no evidence for maternal transmission of bacterial endosymbionts controlling parthenogenesis. Thelytokous parthenogenesis of honeybee workers appears to be a classical qualitative trait, because we did not observe mixed parthenogenesis (amphitoky), which might be expected in the case of multi-locus inheritance.     

Genetic variability of arrhenotokous and thelytokous Venturia canescens (Hymenoptera)

The ichneumonid wasp Venturia canescens (Hymenoptera) has been studied extensively for foraging behaviour and population dynamics of sexually (arrhenotokous) and parthenogenetically (thelytokous) reproducing individuals. Here we report the development of a set of microsatellite markers for V.canescens and use them to show that arrhenotokous individuals have more genetic variability than thelytokous ones, which are even homozygous for all tested loci. Crosses between arrhenotokous individuals suggested one marker, Vcan071, to be linked with the Complementary Sex Determiner (CSD) locus and one, Vcan109, with the Virus Like Protein (vlp-p40) locus. The genome size of V. canescens was estimated to be 274-279?Mb. We discuss how both reproductive modes can give rise to the observed genetic variability and how the new markers can be used for future genetic studies of V. canescens.     

The biosystematic relations between a thelytokous and an arrhenotokous form ofAphytis mytilaspidis (Le Baron) [Hymenoptera: aphelinidae] I. The Reproductive relations

The reproductive relations and the probability of hybridization between an arrhenotokous and a thelytokous form ofA. mytilaspidis were investigated so as to determine the significance of thelytoky in biosystematic studies and its value as a taxonomic character. Gene markers were utilized to detect and follow the various phases of the introgressive process between the arrhenotokous and thelytokous forms. The two forms show only a partiel sexual isolation, the major barrier is prezygotic, arrhenotokous males are much less efficient in recognizing the thelytokous female and copulating with them. A thelytokous female, once inseminated, will utilize the sperm as efficiently as the arrhenotokous female. Viable and fertile hybrids are produced and when conditions are favorable and a suitable host is present the introgressive process will result in a hybrid swarm. The hybrids acquire traits carried by both ancestral stocks.