REDUCED MALE ALLOCATION IN THE PARTHENOGENETIC HERMAPHRODITE DUGESIA POLYCHROA

Parthenogenetic lineages that arise in a hermaphroditic, sexual population will inherit the male function from their sexual progenitors. Natural selection then acts to reduce male allocation of the parthenogens, freeing resources presumably for the female function. Depending on age and the available genetic variation, one therefore expects to find reduced male allocation in naturally occurring parthenogenetic lineages. We investigated the allocation to sperm production in the hermaphroditic flatworm Dugesia polychroa in three lakes containing a sexual (S), a (pseudogamous) parthenogenetic (P), and a mixed sexual-parthenogenetic population (M). Parthenogenetic lineages from M were assumed to be relatively young due to recurrent origins from the coexisting sexuals, whereas those from P were assumed to be older on biogeographical grounds. As predicted, we found drastically reduced sperm production in parthenogens compared to sexuals, even in the parthenogenetic lineages from M, which may be younger. M parthenogens did not have more testes, but produced more sperm than individuals from the purely parthenogenetic population (P). However, the latter result could not be reproduced with laboratory-raised animals and therefore may be a consequence of different ecological conditions in the different lakes, for example, differences in mating rates. To study the behavioral component of male allocation, copulation frequencies were recorded for sexuals from M and for parthenogens from P. Compared to the drastic reduction in sperm production, copulation frequency was less reduced in parthenogens. This may be a consequence of allosperm limitation in pseudogamous parthenogenetic populations.     

Population models of sperm-dependent parthenogenesis

Organisms that reproduce by sperm-dependent parthenogenesis are asexual clones that require sperm of a sexual host to initiate egg production, without the genome of the sperm contributing genetic information to the zygote. Although sperm-dependent parthenogenesis has some of the disadvantages of sex (requiring a mate) without the counterbalancing advantages (mixing of parental genotypes), it appears amongst a wide variety of species. We develop initial models for the density-dependent dynamics of animal populations with sperm-dependent parthenogenesis (pseudogamy or gynogenesis), based on the known biology of the common Enchytraeid worm Lumbricillus lineatus. Its sperm-dependent parthenogenetic populations are reproductive parasites of the hermaphrodite sexual form. Our logistic models reveal two alternative requirements for coexistence at density-dependent equilibria: (i) If the two forms differ in competitive ability, the form with the lower intrinsic birth rate must be compensated by a more than proportionately lower competitive impact from the other, relative to intraspecific competition, (ii) If the two forms differ in their intrinsic capacity to exploit resources, the sperm-dependent parthenogen must be superior in this respect and must have a lower intrinsic birth rate. In general for crowded environments we expect a sperm-dependent parthenogen to compete strongly for limiting resources with the sexual sibling species. Its competitive impact is likely to be weakened by its genetic uniformity, however, and this may suffice to cancel any advantage of higher intrinsic growth rate obtained from reproductive investment only in egg production. We discuss likely thresholds of coexistence for other sperm-dependent parthenogens. The fish Poeciliopsis monacha-lucida likewise obtains an intrinsic growth advantage from reduced investment in male gametes, and so its persistence is likely to depend on it being a poor competitor. The planarian Schmidtea polychroa obtains no such intrinsic benefit because it produces fertile sperm, and its persistence may depend on superior resource exploitation.     

Spatial and ecological overlap between coexisting sexual and parthenogenetic Schmidtea polychroa (Tricladida; Platyhelminthes)

Theoretical models on the costs and benefits of sexual reproduction usually assume that sexual and parthenogenetic individuals coexist and are identical, except for their mode of reproduction. Empirical studies, however, show that conspecific sexuals and parthenogens can differ in ecological preferences and geographical distribution, which complicates the investigation of the costs and benefits of sex. The freshwater planarian Schmidtea polychroa exists in a sexual and a sperm-dependent, parthenogenetic form. The latter produce fertile sperm and mate, but received sperm is used only to induce parthenogenetic embryo development. We compared the spatial and ecological distribution between forms within a lake from which both had been reported. Forty samples showed large differences in the relative frequencies of sexuals and parthenogens. Nineteen samples contained both biotypes. All but one of the 13 ecological parameters that we measured, could not explain a significant part of the variance in relative abundance of each type. Only leech abundance had a significant, negative effect on the presence of sexual individuals. The causes of this effect remained unclear. We also estimated the amount of genetic isolation between sites and between reproductive modes, using body coloration as a genetic marker. Large differences were found between sites, suggesting isolation of local populations by migration barriers. There were smaller differences between sexuals and parthenogens within sites, suggesting that genetic exchange between biotypes may be limited. We conclude that there appears to be weak niche differentiation between sexuals and parthenogens in Lago di Caldonazzo in late summer. Fluctuations in relative frequency appears to be a consequence of low dispersal between local populations and stochastic effects within them. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals

In theory, parthenogenetic lineages have low evolutionary potential because they inexorably accumulate deleterious mutations and do not generate much genotypic diversity. As a result, most parthenogenetic taxa occupy the terminal nodes of phylogenetic trees. The rate and mode of development of parthenogenesis are important factors to consider when assessing its costs and benefits since they determine both the level of genetic diversity and the ecological adaptability of the resulting lineages. The origin of parthenogenesis is polyphyletic in many taxa, suggesting that genetic systems maintaining sexuality are often labile. In addition, the loss of sex may be achieved in several ways, leading to parthenogenetic lineages with distinct genetic profiles. This could then influence not only the fate of such lineages in the long term, but also the outcome of competition with their sexual counterparts in the short term. In this paper, we review the possible evolutionary routes to parthenogenesis based on a survey of the phylogenetic relationships between sexual and parthenogenetic lineages in a broad range of animals. We also examine the different mechanisms by which parthenogenetic lineages could arise, and discuss the influence of these mechanisms on both the genetic properties and the ecological life styles of the resulting lineages.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 151–163.     

Male mate choice and sperm allocation in a sexual/asexual mating complex of Poecilia (Poeciliidae, Teleostei)

Male mate choice is critical for understanding the evolution and maintenance of sexual/asexual mating complexes involving sperm-dependent, gynogenetic species. Amazon mollies (Poecilia formosa) require sperm to trigger embryogenesis, but the males (e.g. Poecilia mexicana) do not contribute genes. Males benefit from mating with Amazon mollies, because such matings make males more attractive to conspecific females, but they might control the cost of such matings by providing less sperm to Amazon mollies. We examined this at the behavioural and sperm levels. P. mexicana males preferred to mate with, and transferred more sperm to conspecific females. However, if males mated with P. formosa, sperm was readily transferred. This underscores the importance of male choice in this system.     

Origin and Genetic Diversity of Diploid Parthenogenetic Artemia in Eurasia

There is wide interest in understanding how genetic diversity is generated and maintained in parthenogenetic lineages, as it will help clarify the debate of the evolution and maintenance of sexual reproduction. There are three mechanisms that can be responsible for the generation of genetic diversity of parthenogenetic lineages: contagious parthenogenesis, repeated hybridization and microorganism infections (e.g. Wolbachia). Brine shrimps of the genus Artemia (Crustacea, Branchiopoda, Anostraca) are a good model system to investigate evolutionary transitions between reproductive systems as they include sexual species and lineages of obligate parthenogenetic populations of different ploidy level, which often co-occur. Diploid parthenogenetic lineages produce occasional fully functional rare males, interspecific hybridization is known to occur, but the mechanisms of origin of asexual lineages are not completely understood. Here we sequenced and analysed fragments of one mitochondrial and two nuclear genes from an extensive set of populations of diploid parthenogenetic Artemia and sexual species from Central and East Asia to investigate the evolutionary origin of diploid parthenogenetic Artemia, and geographic origin of the parental taxa. Our results indicate that there are at least two, possibly three independent and recent maternal origins of parthenogenetic lineages, related to A. urmiana and Artemia sp. from Kazakhstan, but that the nuclear genes are very closely related in all the sexual species and parthenogegetic lineages except for A. sinica, who presumable took no part on the origin of diploid parthenogenetic strains. Our data cannot rule out either hybridization between any of the very closely related Asiatic sexual species or rare events of contagious parthenogenesis via rare males as the contributing mechanisms to the generation of genetic diversity in diploid parthenogenetic Artemia lineages.     

Sex and clonality in the little fire ant

Reproduction systems are controlling the creation of new genetic variants as well as how natural selection can operate on these variants. Therefore, they had historically been one of the main foci of evolutionary biology studies. The little fire ant, Wasmannia auropunctata, has been found to display an extraordinary reproduction system, in which both males and female queens are produced clonally. So far, native sexual populations of W. auropunctata have not been identified. Our goals were to identify such sexual populations and investigate the origins of female parthenogenesis and male clonality. Using mitochondrial DNA and microsatellite markers in 17 native populations, we found that traditional sexual populations occurred in W. auropunctata and are likely the recent source of neighboring clonal populations. Queen parthenogenesis has probably evolved several times through mutational events. Male clonality is tightly linked to queen parthenogenesis and thus appears to be female controlled. Its origin could be accounted for by 2 mutually exclusive hypotheses: either by the expected coevolution of the 2 sexes (i.e., a variant of the maternal genome elimination hypothesis) or by a shared mechanistic origin (i.e., by the production of anucleate ovules by parthenogenetic queens). Our results also show that W. auropunctata males and females do not form separate evolutionary units and are unlikely to be engaged in an all-out battle of sexes. This work opens up new perspectives for studies on the adaptive significance and evolutionary stability of mixed sexual and clonal reproduction systems in living organisms.     

Population genetics and ecology of Artemia: Insights into parthenogenetic reproduction

The relative advantages of sexual and parthenogenetic reproduction have long interested biologists and remain a central issue in ecological and evolutionary studies. Recent data on brine shrimp (Artemia) indicate that extensive ecological and genetic divergence occurs in an obligately parthenogenetic lineage. This challenges the belief that parthenogenetic lineages are evolutionary 'dead ends' and that extensive divergence is necessarily linked to recent recruitment from sexual ancestors. The molecular evidence suggests that parthenogenesis in Artemia is relatively ancient, with a single asexual lineage branching from an Old World sexual ancestor approximately five million years ago. Automictic recombination (which can occur in diploid but not polyploid parthenogenetic brine shrimp) appears to play a central role in the long-term maintenance of the parthenogenetic lineage.     

Obligate asex in a rotifer and the role of sexual signals

Transitions to asexuality have occurred in many animals and plants, yet the biological mechanisms causing such transitions have often remained unclear. Cyclical parthenogens, such as cladocerans, rotifers or aphids often give rise to obligate asexual lineages. In many rotifers, chemical signals that accumulate during population crowding trigger the induction of sexual stages. In this study, I tested two hypotheses on the origin of obligate parthenogenesis in the rotifer Brachionus calyciflorus: (i) that obligate parthenogens have lost the responsiveness to the sexual signal; and (ii) that obligate parthenogens have lost the ability to produce the sexual signal. Pairwise cross-induction assays among three obligate parthenogenetic strains and two cyclically parthenogenetic (sexual) strains were used to test these hypotheses. I found that obligate parthenogens can induce sexual reproduction in sexual strains, but not vice versa. This demonstrates that obligate parthenogens do still produce the sexual signal, but have lost responsiveness to that signal.     

Making it on their own: sperm-dependent hybrid fishes (Cobitis) switch the sexual hosts and expand beyond the ranges of their original sperm donors

Interspecific hybridization may result in asexual hybrid lineages that reproduce via parthenogenesis. Contrary to true parthenogens, sperm-dependent asexuals (gynogens and hybridogens) are restricted to the range of bisexual species, generally the parental taxa, by their need for a sperm donor. It has been documented that asexual lineages may rarely use sperm from a non-parental species or even switch a host. The available literature reports do not allow distinguishing, between whether such host switches arise by the expansion of asexuals out of their parental's range (and into that of another's) or by the local extinction of a parental population followed by a host switch. The present study combines new and previously collected data on the distribution and history of gynogenetic spined loaches (Cobitis) of hybrid origin. We identified at least three clonal lineages that have independently switched their sperm dependency to different non-parental Cobitis species, and in cases incorporated their genomes. Our current knowledge of European Cobitis species and their hybrids suggests that this pattern most probably results from the expansion of gynogenetic lineages into new areas. Such expansion was independent of the original parental species. This suggests that sperm dependence is not as restrictive to geographical expansion when compared with true parthenogenesis as previously thought.     

Increased capacity for sustained locomotion at low temperature in parthenogenetic geckos of hybrid origin

The evolution of parthenogenesis is typically associated with hybridization and polyploidy. These correlates of parthenogenesis may have important physiological consequences that need be taken into account in understanding the relative merits of sexual and parthenogenetic reproduction. We compared the thermal sensitivity of aerobically sustained locomotion in hybrid/triploid parthenogenetic races of the gecko Heteronotia binoei and their diploid sexual progenitors. Endurance times at low temperature (10 degrees , 12.5 degrees , and 15 degrees C, 0.05 km h(-1)) were significantly greater in parthenogenetic females than in sexual females. Comparison of oxygen consumption rates during sustained locomotion at increasing speeds (0.05, 0.10, 0.15, 0.20, 0.25, and 0.30 km h(-1), 25 degrees C) indicated that parthenogenetic lizards have higher maximum oxygen consumption rates and maximum aerobic speeds than do female sexual geckos. In addition, parthenogenetic geckos showed greater levels of voluntary activity at 15 degrees C than did sexual geckos, although this pattern appears strongest in comparison to male sexual forms. Parthenogenetic lineages of Heteronotia thus have an advantage over sexual lineages in being capable of greater aerobic activity. This result is opposite of that found in prior studies of parthenogenetic teiid lizards (genus Cnemidophorus) and highlights the idiosyncratic nature of phenotypic evolution in parthenogens of hybrid origin.     

Facultative symbiont infections affect aphid reproduction

Some bacterial symbionts alter their hosts reproduction through various mechanisms that enhance their transmission in the host population. In addition to its obligatory symbiont Buchnera aphidicola, the pea aphid Acyrthosiphon pisum harbors several facultative symbionts influencing several aspects of host ecology. Aphids reproduce by cyclical parthenogenesis whereby clonal and sexual reproduction alternate within the annual life cycle. Many species, including the pea aphid, also show variation in their reproductive mode at the population level, with some lineages reproducing by cyclical parthenogenesis and others by permanent parthenogenesis. While the role of facultative symbionts has been well studied during the parthenogenetic phase of their aphid hosts, very little is known on their possible influence during the sexual phase. Here we investigated whether facultative symbionts modulate the capacity to produce sexual forms in various genetic backgrounds of the pea aphid with controlled symbiont composition and also in different aphid genotypes from natural populations with previously characterized infection status and reproductive mode. We found that most facultative symbionts exhibited detrimental effects on their hosts fitness under sex-inducing conditions in comparison with the reference lines. We also showed that the loss of sexual phase in permanently parthenogenetic lineages of A. pisum was not explained by facultative symbionts. Finally, we demonstrated that Spiroplasma infection annihilated the production of males in the host progeny by inducing a male-killing phenotype, an unexpected result for organisms such as aphids that reproduce primarily through clonal reproduction.     

The Persistence of Facultative Parthenogenesis in Drosophila albomicans

Parthenogenesis has evolved independently in more than 10 Drosophila species. Most cases are tychoparthenogenesis, which is occasional or accidental parthenogenesis in normally bisexual species with a low hatching rate of eggs produced by virgin females; this form is presumed to be an early stage of parthenogenesis. To address how parthenogenesis and sexual reproduction coexist in Drosophila populations, we investigated several reproductive traits, including the fertility, parthenogenetic capability, diploidization mechanisms, and mating propensity of parthenogenetic D. albomicans. The fertility of mated parthenogenetic females was significantly higher than that of virgin females. The mated females could still produce parthenogenetic offspring but predominantly produced offspring by sexual reproduction. Both mated parthenogenetic females and their parthenogenetic-sexual descendants were capable of parthenogenesis. The alleles responsible for parthenogenesis can be propagated through both parthenogenesis and sexual reproduction. As diploidy is restored predominantly by gamete duplication, heterozygosity would be very low in parthenogenetic individuals. Hence, genetic variation in parthenogenetic genomes would result from sexual reproduction. The mating propensity of females after more than 20 years of isolation from males was decreased. If mutations reducing mating propensities could occur under male-limited conditions in natural populations, decreased mating propensity might accelerate tychoparthenogenesis through a positive feedback mechanism. This process provides an opportunity for the evolution of obligate parthenogenesis. Therefore, the persistence of facultative parthenogenesis may be an adaptive reproductive strategy in Drosophila when a few founders colonize a new niche or when small populations are distributed at the edge of a species'' range, consistent with models of geographical parthenogenesis.     

Patterns and mechanisms in instances of endosymbiont‐induced parthenogenesis

Female‐producing parthenogenesis can be induced by endosymbionts that increase their transmission by manipulating host reproduction. Our literature survey indicates that such endosymbiont‐induced parthenogenesis is known or suspected in 124 host species from seven different arthropod taxa, with Wolbachia as the most frequent endosymbiont (in 56–75% of host species). Most host species (81%, 100 out of 124) are characterized by haplo‐diploid sex determination, but a strong ascertainment bias likely underestimates the frequency of endosymbiont‐induced parthenogenesis in hosts with other sex determination systems. In at least one taxon, hymenopterans, endosymbionts are a significant driver of transitions from sexual to parthenogenetic reproduction, with one‐third of lineages being parthenogenetic as a consequence of endosymbiont infection. Endosymbiont‐induced parthenogenesis appears to facilitate the maintenance of reproductive polymorphism: at least 50% of species comprise both sexual (uninfected) and parthenogenetic (infected) strains. These strains feature distribution differences similar to the ones documented for lineages with genetically determined parthenogenesis, with endosymbiont‐induced parthenogens occurring at higher latitudes than their sexual relatives. Finally, although gamete duplication is often considered as the main mechanism for endosymbiont‐induced parthenogenesis, it underlies parthenogenesis in only half of the host species studied thus far. We point out caveats in the methods used to test for endosymbiont‐induced parthenogenesis and suggest specific approaches that allow for firm conclusions about the involvement of endosymbionts in the origin of parthenogenesis.     

The effect of rearing temperature on body shape and meristic characters in zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>) juveniles

Considering its immediate costs of producing dispensable males, the maintenance of sexual reproduction is a major paradox in evolutionary biology. Asexual lineages that do not face such costs theoretically should replace sexuals over time. Nonetheless, several systems are known in which closely related sexual and asexual lineages stably coexist. In the present study, we studied a sexual/asexual mating complex of a sperm-dependent parthenogenetic fish (amazon molly, Poecilia formosa) and its sexual congeners, the sailfin molly P. latipinna and the Atlantic molly P. mexicana. We asked whether differences in feeding behavior could contribute to their stable coexistence. We conducted a laboratory experiment to compare feeding efficiencies and also measured the competitive abilities between the two reproductive forms. Additionally, we measured gut fullness of fishes caught in natural habitats. Contrary to our predictions, we could not find P. formosa to be less efficient in feeding. We argue that food competition in mollies plays a minor role in mediating coexistence between closely related asexual and sexual mollies.     

Evolutionary implications of developmental instability in parthenogenetic drosophila mercatorum. I. Comparison of several strains with different genotypes

Natural populations of sexually reproducing Drosophila mercatorum are capable of a very low rate of parthenogenesis, but this mode of reproduction has apparently never characterized an entirely asexual population in this species. The high abortion rate observed in laboratory parthenogenetic lines suggests that developmental constraints may cause the failure of this trait to spread in nature. To investigate the basis of this developmental instability and how it may affect the evolution of parthenogenesis in natural populations, early embryonic development was compared between one sexual and four parthenogenetic laboratory strains of D. mercatorum. There is a large amount of variation within a given parthenogenetic strain, suggesting that parthenogenesis is associated with a general breakdown of developmental stability. There is relatively little variation among different parthenogenetic strains, suggesting that most abortions are due to a feature inherent to parthenogenetic reproduction rather than a feature of a particular genome. Likewise, there is little variation between parthenogenetic and sexual strains in the causes of abortions, suggesting that the developmental problems encountered by parthenogenetic lineages are not unique to parthenogens. Thus, the failure of parthenogenesis to spread within D. mercatorum can be attributed to no particular developmental constraint per se operating after the initiation of embryogenesis. However, the overall increase in all developmental problems that occurs with the transition from sexual to parthenogenetic development suggests that the high degree of developmental instability associated with parthenogenesis may be considered a developmental constraint in its own right.     

Random genetic drift during cyclical ameiotic parthenogenesis

The classical models of population genetics assume sexual reproduction and do not apply to organisms in which parthenogenetic reproduction is alternated with sexual recombination. Under cyclic parthenogenesis, variation in rates or frequencies of parthenogenetic reproduction among clones produces selection that is independent of processes occurring in the sexual phases.In this paper I examine how selection during cyclic parthenogenesis influences random genetic drift and leads to a loss of variance among clones. To illustrate these effects, computer simulations are presented demonstrating the response of effective clone number and equilibrium clone diversity to selection and mutation.     

Life-history changes that accompany the transition from sexual to parthenogenetic reproduction in Drosophila mercatorum

In spite of the predicted genetic and ecological costs of sex, most natural populations maintain sexual reproduction, even those capable of facultative parthenogenesis. Unfertilized eggs from natural populations of Drosophila mercatorum occasionally develop into viable adults, but obligately parthenogenetic populations are unknown in this species. To evaluate the microevolutionary forces that both favor and constrain the evolution of parthenogenesis in D. mercatorum, we have measured parthenogenetic rates across a natural, sexually reproducing population and characterized the life-history changes that accompany the transition from sexual to parthenogenetic reproduction in laboratory strains. A highly significant difference in parthenogenetic rate was found between two populations in close geographic proximity, with increased rate found with lower population density. Laboratory strains of parthenogenetic females suffered increased mortality and reduced egg viability relative to their virgin counterparts from a sexual strain. Lifetime egg production was similar across all strains, but a shift in peak egg production to an earlier age also occurred. The combination of these life-history traits resulted in a higher net reproductive value for sexual females, but because they also had a longer generation time, intrinsic rate of increase was not as dramatically different from parthenogenetic females. In environments with high early mortality, there may be no fitness disadvantage to parthenogenesis, but the predicted ecological advantage of a twofold increase in intrinsic rate of increase was not realized. These results support the theory of Stalker (1956) that parthenogenesis is favored in environments in which sexual reproduction is difficult or impossible.     

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.     

Multiple direct transitions from sexual reproduction to apomictic parthenogenesis in Timema stick insects

Transitions from sexual reproduction to parthenogenesis may occur along multiple evolutionary pathways and involve various cytological mechanisms to produce diploid eggs. Here, we investigate routes to parthenogenesis in Timema stick insects, a genus comprising five obligate parthenogens. By combining information from microsatellites and karyotypes with a previously published mitochondrial phylogeny, we show that all five parthenogens likely evolved spontaneously from sexually reproducing species, and that the sexual ancestor of one of the five parthenogens was probably of hybrid origin. The complete maintenance of heterozygosity between generations in the five parthenogens strongly suggests that eggs are produced by apomixis. Virgin females of the sexual species were also able to produce parthenogenetic offspring, but these females produced eggs by automixis. High heterozygosity levels stemming from conserved ancestral alleles in the parthenogens suggest, however, that automixis has not generated the current parthenogenetic Timema lineages but that apomixis appeared abruptly in several sexual species. A direct transition from sexual reproduction to (at least functional) apomixis results in a relatively high level of allelic diversity and high efficiency for parthenogenesis. Because both of these traits should positively affect the demographic success of asexual lineages, spontaneous apomixis may have contributed to the origin and maintenance of asexuality in Timema .