Sexual and asexual colony foundation and the mechanism of facultative parthenogenesis in the termite <Emphasis Type="Italic">Reticulitermes speratus</Emphasis> (Isoptera,Rhinotermitidae)

Summary. Facultative parthenogenesis has great adaptive significance, especially with regard to low pairing efficiency. In the termite Reticulitermes speratus, females that fail to mate with males reproduce parthenogenetically and found colonies cooperatively with partner females or even alone. Comparison of colony foundation success at 400 days between colonies founded by single females (F), female-female pairs (FF), and male-female pairs (FM) showed that female-female cooperation promoted colony survivorship over monogamous foundation. We report here for the first time the mode of parthenogenesis in Isoptera. Combining chromosome observations and genetic analysis using microsatellites, we show that the mode of parthenogenesis is diploid thelytoky and that the restoration of ploidy is most likely accomplished by terminal fusion. Parthenogens show a higher mortality and a longer egg-development time than sexually produced offspring, probably due to reduced heterozygosity. In addition Wolbachia bacteria were detected in R. speratus. However, since Wolbachia was also detected in non-parthenogenetic R. flavipes, it is unlikely that Wolbachia is the cause of parthenogenesis in R. speratus.Received 30 October 2003; revised 4 March 2004; accepted 17 March 2004.     

Comparison of colony foundation success between sexual pairs and female asexual units in the termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

In termites, a male and a female usually found a colony cooperatively. However, pairing efficiency tends to be low in Reticulitermes speratus because of a limited mate-searching range, the female-biased sex ratio, and a relatively low calling ability. Females that fail to pair with males found colonies either in female–female pairs or even alone. In the laboratory, we examined colony foundation by single females (F), female–female pairs (FF), and normal male–female pairs (FM). The time until colony foundation (when termites began excavating wood baits) differed significantly among the unit types. Time until excavation was much longer for single females than for FF and FM units, which reflects the relative success of colony foundation. The survival rate of single females was also significantly lower than that of FF- and FM-unit females, although there was no difference between FF and FM units. This result demonstrates that cooperation, even female–female, promotes female survivorship. Nevertheless, the number of progeny per female was significantly lower in FF units than in FM units, possibly because females of FF units must share reproductive output. These results lead us to the conclusion that a normal monogamous pair is the best unit for colony foundation. Nevertheless, females alone can establish colonies by parthenogenesis, and even female–female cooperation promotes colony foundation success if pairing with males is not possible. Considering the functional decision for females in F and FF units of how much time to spend searching for a male mate, we believe that these facultative pathways of colony foundation by parthenogenesis have adaptive significance. Received: November 30, 2000 / Accepted: March 6, 2001     

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.     

Cooperative colony foundation by termite female pairs: altruism for survivorship in incipient colonies

To determine the costs and benefits of queen association in termites we examined for the first time female-female interactions in colonies founded by two unmated females. In the termite Reticulitermes speratus, females that fail to pair with males found colonies cooperatively with partner females and reproduce by parthenogenesis. We analysed the relationship between queen dominance and initial size ranking in two-queen colonies from the viewpoint of first worker brood production and weight gain. To assign parentage to offspring of two-queen colonies we used mtDNA restriction fragment length polymorphism (RFLP); the results suggested that the two queens produced first worker brood equally throughout colony foundation. Furthermore, initially smaller females gained significantly more weight than initially larger females. This may have resulted from altruistic behaviour of the larger females towards the smaller ones. A simple mathematical model, which considered resource allocation and survivorship, could explain why the larger females behave altruistically towards the smaller females. We also examined the responses of females when more than two females were placed in a petri dish in the presence or absence of a male. If a partner male was present, only one female survived in the colony. In the absence of a partner male, two females, but never more than two, founded a colony cooperatively. These results show that females need a partner to found, and retain, a colony.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Patterns of neotenic differentiation in a subterranean termite, Reticulitermes speratus (Isoptera: Rhinotermitidae)

Plasticity in the caste developmental pathway is a remarkable characteristic of termite societies. In Reticulitermes, two types of neotenic reproductive, nymphoids and ergatoids, may differentiate from nymphs and workers and take over reproduction in the colony after the death of the original primary reproductive pair. We examined the dynamics of newly differentiated nymphoids and ergatoids in experimentally orphaned laboratory colonies of R. speratus with different caste compositions. The period required for differentiation of nymphoids was shorter than that for differentiation of ergatoids. The sex ratio of neotenics was strongly female‐biased, particularly in ergatoids. The results suggested that the number of differentiated ergatoids was restricted by the existence of nymphs or nymphoids in a colony. Workers were assumed to kill most newly differentiated neotenics. Attack reflecting conflict between colony members is probably an important mechanism to control neotenic emergence.     

First molecular genetic evidence for automictic parthenogenesis in cockroaches

Parthenogenesis is an asexual mode of reproduction that plays an important role in the evolution of sex, sociality, and reproduction strategies in insects. Some species of cockroach exhibit thelytoky, a type of parthenogenesis in which female offspring are produced without fertilization. However, the cytological and genetic mecha? nisms of parthenogenesis in cockroaches are not well understood. Here we provide the first molecular genetic evidence that cockroaches can reproduce through automixis. Using the American cockroach Periplaneta aniericana, we performed microsatellite analysis to investigate the genetic relationship between parthenogenetically produced nymphs and the parent virgin females, and found that all parthenogenetic offspring were homozygous for autosomal microsatellite markers, whereas the female parents were heterozygous. In addition, flow cytometry analysis revealed that the parthenogenetic offspring were diploid. Taken together, our results demonstrate that P. americana exhibits automixis-type thelytoky, in which diploidy is restored by gamete duplication or terminal fusion. These findings highlight the unique reproduction strategies of cockroaches, which are more varied than was previously recognized.     

Sex at the margins: parthenogenesis vs. facultative and obligate sex in a Neotropical ant

Geographic parthenogenesis is a distribution pattern, in which parthenogenetic populations tend to live in marginal habitats, at higher latitudes and altitudes and island‐like habitats compared with the sexual forms. The facultatively parthenogenetic ant Platythyrea punctata is thought to exhibit this general pattern throughout its wide range in Central America and the Caribbean Islands. Workers of P. punctata from the Caribbean produce diploid female offspring from unfertilized eggs by thelytokous parthenogenesis, and mated females and males are rare. In contrast, workers in one colony from Costa Rica were incapable of thelytoky; instead mated workers produced all female offspring. Because sample sizes were very low in former studies, we here use microsatellite markers and explicit tests of thelytoky to examine the population genetic structure of ancestral and derived populations of P. punctata throughout the Caribbean and Central America. Populations from the Caribbean islands were fully capable of parthenogenesis, and population genetic signatures indicate that this is the predominant mode of reproduction, although males are occasionally produced. In contrast, the northernmost population on the mainland (Texas) showed signatures of sexual reproduction, and individuals were incapable of reproduction by thelytoky. Contrary to expectations from a geographic parthenogenesis distribution pattern, most parts of the mainland populations were found to be facultatively thelytokous, with population genetic signatures of both sexual and parthenogenetic reproduction.     

Geographic parthenogenesis and the common tea‐tree stick insect of New Zealand

Worldwide, parthenogenetic reproduction has evolved many times in the stick insects (Phasmatidae). Many parthenogenetic stick insects show the distribution pattern known as geographic parthenogenesis, in that they occupy habitats that are at higher altitude or latitude compared with their sexual relatives. Although it is often assumed that, in the short term, parthenogenetic populations will have a reproductive advantage over sexual populations; this is not necessarily the case. We present data on the distribution and evolutionary relationships of sexual and asexual populations of the New Zealand stick insect, Clitarchus hookeri. Males are common in the northern half of the species’ range but rare or absent elsewhere, and we found that most C. hookeri from putative‐parthenogenetic populations share a common ancestor. Female stick insects from bisexual populations of Clitarchus hookeri are capable of parthenogenetic reproduction, but those insects from putative‐parthenogenetic populations produced few offspring via sexual reproduction when males were available. We found similar fertility (hatching success) in mated and virgin females. Mated females produce equal numbers of male and female offspring, with most hatching about 9–16 weeks after laying. In contrast, most eggs from unmated females took longer to hatch (21–23 weeks), and most offspring were female. It appears that all C. hookeri females are capable of parthenogenetic reproduction, and thus could benefit from the numerical advantage this yields. Nevertheless, our phylogeographic evidence shows that the majority of all‐female populations over a wide geographic area originate from a single loss of sexual reproduction.     

Fitness of alternative modes of reproduction: developmental constraints and the evolutionary maintenance of sex

Parthenogenesis, including facultative parthenogenesis, is common among orthopteroid insects. We investigated the fitness associated with sexual and asexual reproduction within a population of the facultatively parthenogenetic cockroach Nauphoeta cinerea. There is significantly reduced fitness for females reproducing parthenogenetically compared to sexually. Fewer than half of all females can reproduce parthenogenetically. In addition, tenfold fewer offspring are produced by parthenogenesis due to reductions in both the number of offspring produced per clutch and the number of clutches produced. Development and brooding of sexually or parthenogenetically produced first instar nymphs does not differ, although the production of the first parthenogenetic clutch is delayed relative to the first sexually produced clutch. The fitness of parthenogens is also lower than the fitness of sexually produced offspring. Parthenogens are less viable than sexually produced offspring even in the benign conditions of the laboratory. Development to adulthood of parthenogens is slower. Fewer parthenogens survive to adulthood and the adult life span of parthenogens is reduced. Individuals produced by parthenogenetic reproduction are unlikely to reproduce parthenogenetically themselves. Finally, parthenogenetically produced females produce fewer offspring by sexual reproduction than do sexually produced females. Since parthenogenetic reproduction is apomictic in N. cinerea and parthenogens are diploid, we suggest that asexual reproduction is developmentally constrained. Once meiosis has evolved, returning to a mitotic mode of reproduction may be difficult. Nauphoeta cinerea offers a system for testing how asexuality is constrained as modes of reproduction can be compared within a facultative parthenogen.     

Appearance of males in a thelytokous strain of Milnesium cf. tardigradum (Tardigrada)

Tardigrades are generally gonochoristic. Many moss-dwelling species propagate by parthenogenesis, but heterogony has not yet been found. Milnesium tardigradum, a carnivorous tardigrade, also has both sexes, but males are usually rare and many populations appear to have only parthenogenetic reproduction. Since 2000, I have maintained a thelytokous strain of Milnesium cf. tardigradum that originated from one female. Individuals of this strain were thought to be all females, but here I report that males have emerged in this strain at a very low frequency. This is the first report of the appearance of males in parthenogenetic tardigrades. On the first pair of legs of some individuals, I observed the modified claws characteristic of males of this species. It is unknown whether these males can actually function in sexual reproduction; however, they might allow some possibility of genetic exchange among clonal populations. No environmental factors that generate males were determined.     

Microsatellite loci in the Japanese subterranean termite,Reticulitermes speratus

Reticulitermes termites have such a cryptic life style and complex colony structure that polymorphic microsatellite markers are desired to investigate their population and colony structures. We successfully isolated seven microsatellite loci in R. speratus , the Japanese subterranean termite, five of which were polymorphic. These polymorphic loci had 2–8 alleles per locus and their observed heterozygosities ranged from 0.059 to 0.438. These five loci were also polymorphic in R . kanmonensis , distributed sympatrically with R. speratus in the Kanmon Region, western Japan; the number of alleles per locus was 2–5, and observed heterozygosity was 0.176–0.625.     

Breeding systems and reproductive strategies in Italian Reticulitermes colonies (Isoptera: Rhinotermitidae)

Biological traits and colony structure are difficult to analyze in subterranean termites owing to their cryptic lifestyle and their often elusive breeding system. However, the use of molecular markers in a population genetics framework allows the investigation of such aspects. We present here the colony genetic structures of 12 samples collected along the Italian peninsula of two Reticulitermes species (the native R. lucifugus and the introduced R. urbis) analyzed through nuclear microsatellite markers. Reproductive strategies and colony breeding systems differ between the two species. Secondary reproductives of R. lucifugus, collected in three colonies, are all females; genotyping comparisons between these females and their nest mate workers clearly indicate the presence of asexual queen succession (AQS) events in this species, as observed in the Japanese R. speratus and in the North-American R. virginicus. Two other R. lucifugus colonies have a mixed family genetic pattern, possibly as the result of colony fusion events: accordingly, relatedness estimates indicate the presence of genetically unrelated workers. On the contrary, all R. urbis colonies have a genetic structure compatible with the presence of multiple secondary reproductives, as expected on the basis of previous analyses. Moreover, neotenics’ sex ratio is balanced and their heterozygosity is comparable to that of nest mate workers, suggesting that AQS is lacking in this taxon. The differences observed in such biological traits between the two species are discussed in the light of their invasive potential.     

Thelytokous parthenogenesis in the exotic dacetine ant Strumigenys rogeri (Hymenoptera: Formicidae) in Taiwan

Only recently has it become clear that several species of eusocial hymenopterans regularly reproduce by thelytokous parthenogenesis, that is, the production of diploid female offspring by unmated females. This phenomenon suggests that parthenogenetic reproduction might be advantageous to organisms under certain environmental conditions. Here the occurrence of asexual reproduction is reported for the first time in the dacetine ant, Strumigenys rogeri, at least for the focal populations in Taiwan. Virgin queens of S. rogeri maintained with several workers produced both workers and young queens from unfertilized eggs under laboratory conditions in as short as 39 days, whereas workers were strictly sterile as no spermatheca was discovered after dissection. Combined with additional evidence (i.e. absence of males in field colonies), queen thelytoky is confirmed. Such a reproductive mode and short development time may jointly help explain the success of this tramp ant species in Taiwan and elsewhere.     

Molecular genetic evidence for parthenogenesis in the Burmese python,Python molurus bivittatus

Parthenogenesis among reptiles is rare. Only a few species have the ability to reproduce asexually. Most of these are obligate parthenogenetic species that consist (almost) entirely of females, which can reproduce solely through parthenogenesis. Rarer are sexual species that only sporadically reproduce through parthenogenesis. A female Python molurus bivittatus (Reptilia, Boidae) from the Artis Zoo, Amsterdam, produced eggs in five consecutive years that contained embryos while she was isolated from males. These eggs might be fertilized with stored sperm, or might be the product of parthenogenesis. Parthenogenesis has not been shown for the Boidae family before. We performed parentship analyses on the snake and seven of her embryos using microsatellites and AFLP. Four microsatellite loci developed for this species combined with three loci developed previously for different snake species revealed too little variation to discriminate between sperm retention and parthenogenesis. With AFLP we were able to confirm that the Artis Zoo female reproduced parthenogenetically. Because the offspring are genetically identical to their mother, whereas in previous studies on sporadic parthenogenesis in snakes a loss of genetic information was reported, we conclude that the meiotic pathways that produce the diploid egg cells are different.     

Genetic variation and asexual reproduction in the facultatively parthenogenetic cockroach Nauphoeta cinerea: implications for the evolution of sex

Asexual reproduction could offer up to a two‐fold fitness advantage over sexual reproduction, yet higher organisms usually reproduce sexually. Even in facultatively parthenogenetic species, where both sexual and asexual reproduction is sometimes possible, asexual reproduction is rare. Thus, the debate over the evolution of sex has focused on ecological and mutation‐elimination advantages of sex. An alternative explanation for the predominance of sex is that it is difficult for an organism to accomplish asexual reproduction once sexual reproduction has evolved. Difficulty in returning to asexuality could reflect developmental or genetic constraints. Here, we investigate the role of genetic factors in limiting asexual reproduction in Nauphoeta cinerea, an African cockroach with facultative parthenogenesis that nearly always reproduces sexually. We show that when N. cinerea females do reproduce asexually, offspring are genetically identical to their mothers. However, asexual reproduction is limited to a nonrandom subset of the genotypes in the population. Only females that have a high level of heterozygosity are capable of parthenogenetic reproduction and there is a strong familial influence on the ability to reproduce parthenogenetically. Although the mechanism by which genetic variation facilitates asexual reproduction is unknown, we suggest that heterosis may facilitate the switch from producing haploid meiotic eggs to diploid, essentially mitotic, eggs.     

The fitness effects of delayed switching to sex in a facultatively asexual insect

Facultative reproductive strategies that incorporate both sexual and parthenogenetic reproduction should be optimal, yet are rarely observed in animals. Resolving this paradox requires an understanding of the economics of facultative asexuality. Recent work suggests that switching from parthenogenesis to sex can be costly and that females can resist mating to avoid switching. However, it remains unclear whether these costs and resistance behaviors are dependent on female age. We addressed these questions in the Cyclone Larry stick insect, Sipyloidea larryi, by pairing females with males (or with females as a control) in early life prior to the start of parthenogenetic reproduction, or in mid‐ or late life after a period of parthenogenetic oviposition. Young females were receptive to mating even though mating in early life caused reduced fecundity. Female resistance to mating increased with age, but reproductive switching in mid‐ or late life did not negatively affect female survival or offspring performance. Overall, mating enhanced female fitness because fertilized eggs had higher hatching success and resulted in more adult offspring than parthenogenetic eggs. However, female fecundity and offspring viability were also enhanced in females paired with other females, suggesting a socially mediated maternal effect. Our results provide little evidence that switching from parthenogenesis to sex at any age is costly for S. larryi females. However, age‐dependent effects of switching on some fitness components and female resistance behaviors suggest the possibility of context‐dependent effects that may only be apparent in natural populations.     

Sexual reproduction of Daphnia pulex in a temporary habitat

Species of Daphnia (Crustacea: Cladocera) typically reproduce by cyclical parthenogenesis, in which a period of all-female parthenogenetic reproduction is followed by sexual reproduction. Sex in Daphnia is determined by the environment, with factors such as temperature, photoperiod and crowding stimulating the production of males and sexual females. Previous studies on Daphnia pulex from temporary pond habitats demonstrated the coexistence of male-producing and non-male-producing (NMP) females, as determined under crowding in the laboratory. A strong genetic component to this sex allocation variation suggested that sex expression in D. pulex is better described as a result of genotype-environment interaction. The present study examined the switch from parthenogenetic to sexual reproduction in two temporary-pond populations of D. pulex. Both populations showed a very early investment in sexual reproduction, independent of population density, by producing males very soon after the populations were reestablished from resting eggs in the early spring. Approximately 40% of the initial broods were male. Additional evidence for gender specialization was obtained by observing the sex of two or three successive broods for 85 individual females. Fifty-eight females produced successive broods of females, 13 females produced successive broods of males and 14 females produced successive broods which included both male and female broods. Females that produced successive female broods under natural conditions included a higher frequency of NMP females compared to a random sample of females, confirming the existence of NMP females. Sexual females were observed in both populations after the first appearence of males, suggesting that the presence of males may stimulate the production of sexual females. For D. pulex populations in a temporary environment, there appears to be an increased emphasis on sexual reproduction and a decreased influence of the environment on sex determination, compared to Daphnia populations in more permanent habitats. Received: 19 February 1996 / Accepted: 20 January 1997     

The Population Genetics of Parthenogenetic Strains of DROSOPHILA MERCATORUM. II. the Capacity for Parthenogenesis in a Natural, Bisexual Population

Drosophila mercatorum is a bisexual species, but certain strains are capable of parthenogenetic reproduction in the laboratory. We investigated the parthenogenetic capacity of the virgin daughters of females captured from a natural, bisexual population in Hawaii. An isozyme survey indicated the natural population is polymorphic at about 50% of its loci, and its individuals heterozygous at 18% of their loci. The predominant mode of parthogenesis in D. mercatorum causes homozygosity for all loci in a single generation. Despite this radical change in genetic state, 23% of the virgin female lines produced adult parthenogenetic progeny, and 16% produced parthenogenetic progeny themselves capable of parthenogenetic reproduction. The parthenogenetic rats as measured by the number of parthenogenetic progeny themselves capable of parthenogenesis divided by the number of eggs laid is arougn 10(-5) for the virgin female lines. We argue that one of the major reasons for this low rate is that very few of the impaternate zygotes have a genotype that can survive and reproduce under the genetic conditions imposed by parthenogenetic reproduction. This intense selective bottleneck can be passed in a single generation if enough unfertilized eggs are laid, and once passed is accompanied by a large (perhaps a thousandfold) increase in the rate of parthenogenesis and by modifications in many phenotypic traits such as morphology and behavior.     

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.     

Evolutionary genetics and ecology of sperm-dependent parthenogenesis

Sperm-dependent (or pseudogamous) forms of parthenogenetic reproduction occur in a wide variety of animals. Inheritance is typically clonal and matroclinous (of female descent), but sperm are needed to initiate normal development. As opposed to true parthenogenesis (i.e., sperm-independent reproduction), pseudogamous parthenogenetic lineages must coexist with a ‘sperm donor’— e.g., males from a conspecific sexual lineage, conspecific hermaphrodites, or males from a closely related sexual species. Such sperm donors do not contribute genetically to the next generation. The parasitic nature of sperm-dependent parthenogenesis raises numerous ecological and evolutionary questions. How do they arise? What factors help stabilize coexistence between the pseudogamous parthenogens and their sperm donors (i.e., ‘sexual hosts’)? Why do males waste sperm on the asexual females? Why does true parthenogenesis not evolve in pseudogamous lineages and free them from their dependency on sperm donors? Does pseudogamous parthenogenesis provide compensatory benefits that outweigh the constraints of sperm-dependence? Herein, we consider some genetic, ecological, and geographical consequences of sperm-dependent parthenogenesis in animals.