Simultaneous hermaphrodites reproducing in pairs self-fertilize some of their eggs: an experimental test of predictions of mixed-mating and Hermaphrodite's Dilemma theory

Theory predicts (1) that mixed-mating systems (i.e. reproduction through both selfing and outcrossing) should usually not evolve and (2) that reproducing simultaneous hermaphrodites should be in a conflict over the preferred sexual role (The Hermaphrodite's Dilemma). In an in vitro system with the endoparasitic cestode Schistocephalus solidus, a simultaneous hermaphrodite, we tested predictions of both the mixed-mating and the Hermaphrodite's Dilemma theory. Using microsatellite markers, we measured the proportion of selfed offspring and the total reproductive output of each worm within pairs varying in mean weight and weight difference. Worms produced more outbred offspring not only with increasing total weight of the pair, but also with decreasing weight difference between the two paired worms. These results suggest: (1) that this parasite species reproduces by mixed-mating, which may be maintained by stochastic density fluctuations in the definitive host and hence unpredictability of self reproduction and (2) reproductive conflict may prevent worm pairs from achieving an optimal intermediate selfing rate.     

Early inbreeding depression in the sexually polymorphic plant Dianthus sylvestris (Caryophyllaceae): Effects of selfing and biparental inbreeding among sex morphs

Predominantly outcrossing plant species are expected to accumulate recessive deleterious mutations, which can be purged when in a homozygous state following selfing. Individuals may vary in their genetic load because of different selfing histories, which could lead to differences in inbreeding depression among families. Lineage-dependent inbreeding depression can appear in gynodioecious species if obligatory outcrossed females are more likely to produce female offspring and if partially selfing hermaphrodites are more likely to produce hermaphrodites. We investigated inbreeding depression at the zygote, seed, and germination stages in the gynomonoecious-gynodioecious Dianthus sylvestris, including pure-sexed plants and a mixed morph. We performed hand-pollinations on 56 plants, belonging to the three morphs, each receiving 2-3 cross treatments (out-, sib- and self-pollination) on multiple flowers. Effects of cross treatments varied among stages and influenced seed provisioning, with sibling competition mainly occurring within outcrossed fruits. We found significant inbreeding depression for seed mass and germination and cumulative early inbreeding depression varied greatly among families. Among sex morphs, we found that females and hermaphrodites differed in biparental inbreeding depression, whereas uniparental was similar for all. Significant inbreeding depression levels may play a role in female maintenance in this species, and individual variation in association with sex-lineages proclivity is discussed.     

SEX‐SPECIFIC INBREEDING DEPRESSION DEPENDS ON THE STRENGTH OF MALE–MALE COMPETITION

Inbreeding depression has become a central theme in evolutionary biology and is considered to be a driving force for the evolution of reproductive morphology, physiology, behavior, and mating systems. Despite the overwhelming body of empirical work on the reproductive consequences of inbreeding, relatively little is known on whether inbreeding depresses male and female fitness to the same extent. However, sex‐specific inbreeding depression has been argued to affect the evolution of selfing rates in simultaneous hermaphrodites and provides a powerful approach to test whether selection is stronger in males than in females, which is predicted to be the consequence of sexual selection. We tested for sex‐specific inbreeding depression in the simultaneously hermaphroditic freshwater snail Physa acuta by comparing the reproductive performance of both sex functions between selfed and outcrossed focal individuals under different levels of male–male competition. We found that inbreeding impaired both male and female reproductive success and that the magnitude of male inbreeding depression exceeded female inbreeding depression when the opportunity for sperm competition was highest. Our study provides the first evidence for sex‐specific inbreeding depression in a hermaphroditic animal and highlights the importance of considering the level of male–male competition when assessing sex differences in inbreeding depression.     

The evolution from females to hermaphrodites results in a sexual conflict over mating in androdioecious nematode worms and clam shrimp

The nematode worm Caenorhabditis elegans and the clam shrimp Eulimnadia texana are two well‐studied androdioecious species consisting mostly of self‐fertilizing hermaphrodites and few males. To understand how androdioecy can evolve, a simple two‐step mathematical model of the evolutionary pathway from a male–female species to a selfing‐hermaphrodite species is constructed. First, the frequency of mutant females capable of facultative self‐fertilization increases if the benefits of reproductive assurance exceed the cost. Second, hermaphrodites become obligate self‐fertilizers if the fitness of selfed offspring exceeds one‐half the fitness of outcrossed offspring. Genetic considerations specific to C. elegans and E. texana show that males may endure as descendants of the ancestral male–female species. These models combined with an extensive literature review suggest a sexual conflict over mating in these androdioecious species: selection favours hermaphrodites that self and males that outcross. The strength of selection on hermaphrodites and males differs, however. Males that fail to outcross suffer a genetic death. Hermaphrodites may never encounter a rare male, and those that do and outcross only bear less fecund offspring. This asymmetric sexual conflict results in an evolutionary stand‐off: rare, but persistent males occasionally fertilize common, but reluctant hermaphrodites. A consequence of this stand‐off may be an increase in the longevity of the androdioecious mating system.     

Effects of reproductive compensation, gamete discounting and reproductive assurance on mating-system diversity in hermaphrodites

Hermaphroditism allows considerable scope for contributing genes to subsequent generations through various mixtures of selfed and outcrossed offspring. The fitness consequences of different family compositions determine the evolutionarily stable mating strategy and depend on the interplay of genetic features, the nature of mating, and factors that govern offspring development. This theoretical article considers the relative contributions of these influences and their interacting effects on mating-system evolution, given a fixed genetic load within a population. Strong inbreeding depression after offspring gain independence selects for exclusive outcrossing, regardless of the intensity of predispersal inbreeding depression, unless insufficient mating limits offspring production. The extent to which selfing evolves under weak postdispersal inbreeding depression depends on predispersal inbreeding depression and the opportunity for resource limitation of offspring production. Mixed selfing and outcrossing is an evolutionarily stable strategy (ESS) if selfed zygotes survive poorly, but selfed offspring survive well, and maternal individuals produce enough "extra" eggs that deaths of unviable outcrossed embryos do not impact offspring production (reproductive compensation). Mixed mating can also be an ESS, despite weak lifetime inbreeding depression, if self-mating reduces the number of male gametes available for outcrossing (male-gamete discounting). Reproductive compensation and male-gamete discounting act largely independently on mating-system evolution. ESS mating systems always involve either complete fertilization or fertilization of enough eggs to induce resource competition among embryos, so although reproductive assurance is adaptive with insufficient mating, it is never an ESS. Our results illustrate the theoretical importance of different constraints on offspring production (availability of male gametes, egg production, and maternal resources) for both the course and outcome of mating-system evolution, whereas unequal competition between selfed and outcrossed embryos has limited effect. These results also underscore the significance of heterogeneity in the nature and intensity of inbreeding depression during the life cycle for the evolution of hermaphrodite mating systems.     

When is it worth being a self-compatible hermaphrodite? Context-dependent effects of self-pollination on female advantage in gynodioecious Silene nutans

In gynodioecious plant species with nuclear‐cytoplasmic sex determination, females and hermaphrodites plants can coexist whenever female have higher seed fitness than hermaphrodites. Although the effect of self fertilization on seed fitness in hermaphrodites has been considered theoretically, this effect is far from intuitive, because it can either increase the relative seed fitness of the females (if it leads hermaphrodites to produce inbred, low quality offspring) or decrease it (if it provides reproductive assurance to hermaphrodites). Hence, empirical investigation is needed to document whether relative seed fitness varies with whether pollen is or is not limiting to seed production. In the current study, we measured fruit set and seed production in both females and hermaphrodites and the selfing rate in hermaphrodites in two experimental patches that differed in sex ratios in the gynodioecious plant Silene nutans. We found an impact of plant gender, patch, and their interaction, with females suffering from stronger pollen limitation when locally frequent. In the most pollen‐limited situation, the selfing rate of hermaphrodites increased and provided hermaphrodites with a type of reproductive assurance that is not available to females. By integrating both the beneficial (reproductive assurance) and costly effects (through inbreeding depression) of self‐pollination, we showed that whether females did or did not exhibit higher seed fitness depended on the degree of pollen limitation on seed production.     

Ovule discounting in an outcrossing, cryptically dioecious tree

Ovule discounting denotes the reduction in the number of ovules available for cross-fertilization due to the interference of inferior pollen. Traditionally, ovule discounting has been discussed solely from the perspective of compromised outcrossing opportunities as a result of selfing, but the principle is more general. Here, we extend its applicability beyond the simple contrast between selfing and outcrossing by showing that, in the cryptically dioecious tree species Fraxinus ornus, ovule discounting through frequent outcrossing with inferior fathers also constitutes a substantial cost of mating. In F. ornus, hermaphrodites produce pollen capable of siring offspring, but these offspring are less viable than those sired by males and are inferred to produce few, if any, surviving progeny. In this paper, we used microsatellite markers to analyze the mating system and paternity in a wild population of F. ornus. We found that the effective number of sires per mother was low (N(ep) = 2.93 to 4.95), and that paternity was correlated among progeny sampled from the same mother, but not among progeny sampled from neighboring mothers. Despite the existence of a local spatial genetic structure (up to 30 m), we found no evidence of biparental inbreeding. There was negligible selfing by hermaphrodites, but they sired approximately one fourth of the seeds produced by other hermaphrodites. Given that these progeny are not inferred to reach reproductive maturity, this constitutes a substantial cost of ovule discounting in the broad sense. We discuss the possible reasons for why hermaphrodites invest resources into inferior pollen.     

Two's company, three's a crowd: experimental evaluation of the evolutionary maintenance of trioecy in Mercurialis annua (Euphorbiaceae)

Trioecy is an uncommon sexual system in which males, females, and hermaphrodites co-occur as three clearly different gender classes. The evolutionary stability of trioecy is unclear, but would depend on factors such as hermaphroditic sex allocation and rates of outcrossing vs. selfing. Here, trioecious populations of Mercurialis annua are described for the first time. We examined the frequencies of females, males and hermaphrodites across ten natural populations and evaluated the association between the frequency of females and plant densities. Previous studies have shown that selfing rates in this species are density-dependent and are reduced in the presence of males, which produce substantially more pollen than hermaphrodites. Accordingly, we examined the evolutionary stability of trioecy using an experiment in which we (a) indirectly manipulated selfing rates by altering plant densities and the frequency of males in a fully factorial manner across 20 experimental plots and (b) examined the effect of these manipulations on the frequency of the three sex phenotypes in the next generation of plants. In the parental generation, we measured the seed and pollen allocations of hermaphrodites and compared them with allocations by unisexual plants. In natural populations, females occurred at higher frequencies in denser patches, a finding consistent with our expectations. Under our experimental conditions, however, no combination of plant densities and male frequencies was associated with increased frequencies of females. Our results suggest that the factors that regulate female frequencies in trioecious populations of M. annua are independent of those regulating male frequencies (density), and that the stable co-existence of all three sex phenotypes within populations is unlikely.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

Postcopulatory sexual selection reduces genetic diversity in experimental populations of Caenorhabditis elegans

Postcopulatory sexual selection affects the evolution of numerous features ranging from mating behavior to seminal fluid toxicity to the size of gametes. In an earlier study of the effect of sperm competition risk on sperm size evolution, experimental populations of the nematode Caenorhabditis elegans were maintained either by outcrossing (sperm competition present) or by selfing (no sperm competition), and after 60 generations, significantly larger sperm had evolved in the outcrossing populations. To determine the effects of this selection on population genetic variation, we assessed genetic diversity in a large number of loci using random amplification of polymorphic DNA-PCR. Nearly 80% of the alleles present in parental strain populations persisted in the 6 experimental populations after the 60 generations and, despite a 2.2-fold difference in expected heterozygosity, the resulting levels of genetic variation were equivalent between the outcrossing and selfing experimental populations. By inference, we conclude that genetic hitchhiking due to sexual selection in the experimental populations dramatically reduced genetic diversity. We use the levels of variation in the selfing populations as a control for the effects of drift, and estimate the strength of sexual selection to be strong in obligatorily outcrossing populations. Although sequential hermaphrodites like C. elegans probably experience little sexual selection in nature, these data suggest that sexual selection can profoundly affect diversity in outcrossing taxa.     

Fitness Effects of Thermal Stress Differ Between Outcrossing and Selfing Populations in <Emphasis Type="Italic">Caenorhabditis elegans</Emphasis>

The maintenance of males and outcrossing is widespread, despite considerable costs of males. By enabling recombination between distinct genotypes, outcrossing may be advantageous during adaptation to novel environments and if so, it should be selected for under environmental challenge. However, a given environmental change may influence fitness of male, female, and hermaphrodite or asexual individuals differently, and hence the relationship between reproductive system and dynamics of adaptation to novel conditions may not be driven solely by the level of outcrossing and recombination. This has important implications for studies investigating the evolution of reproductive modes in the context of environmental changes, and for the extent to which their findings can be generalized. Here, we use Caenorhabditis elegans—a free-living nematode species in which hermaphrodites (capable of selfing but not cross-fertilizing each other) coexist with males (capable of fertilizing hermaphrodites)—to investigate the response of wild type as well as obligatorily outcrossing and obligatorily selfing lines to stressfully increased ambient temperature. We found that thermal stress affects fitness of outcrossers much more drastically than that of selfers. This shows that apart from the potential for recombination, the selective pressures imposed by the same environmental change can differ between populations expressing different reproductive systems and affect their adaptive potential.     

Epigenetic regulation of sex ratios may explain natural variation in self-fertilization rates

Self-fertilization (selfing) favours reproductive success when mate availability is low, but renders populations more vulnerable to environmental change by reducing genetic variability. A mixed-breeding strategy (alternating selfing and outcrossing) may allow species to balance these needs, but requires a system for regulating sexual identity. We explored the role of DNA methylation as a regulatory system for sex-ratio modulation in the mixed-mating fish Kryptolebias marmoratus. We found a significant interaction between sexual identity (male or hermaphrodite), temperature and methylation patterns when two selfing lines were exposed to different temperatures during development. We also identified several genes differentially methylated in males and hermaphrodites that represent candidates for the temperature-mediated sex regulation in K. marmoratus. We conclude that an epigenetic mechanism regulated by temperature modulates sexual identity in this selfing species, providing a potentially widespread mechanism by which environmental change may influence selfing rates. We also suggest that K. marmoratus, with naturally inbred populations, represents a good vertebrate model for epigenetic studies.     

Pollinator service affects quantity but not quality of offspring in a widespread New Zealand endemic tree species

Human-mediated introductions of predatory mammals to New Zealand have led to massive bird extinctions and population declines. This reduction in bird abundance could have a variety of negative effects on the plants they pollinate, and this has led to recent management efforts to eliminate predatory mammals in some areas. In this study, we utilized variation in management efforts to determine the effects of pollinator service on progeny quantity and quality of an endemic New Zealand tree species, Fuchsia excorticata. Due to its gynodioecious breeding system (separate hermaphroditic and female plants), we were able to estimate effective pollination by scoring pollen deposition on females as a proxy for pollinator service, which we used to examine the effect of pollinator service on both the quantity of fruits produced and the quality of fruits, as measured by the selfing rate of hermaphrodites. We found that increased pollinator service increased progeny quantity, with a stronger effect in females, but had no effect on the quality via selfing rates. The combined effects of pollen limitation and loss of seeds due to inbreeding depression led to hermaphrodites having an estimated 24–80% reduction in the number of offspring reaching reproductive age, even in the highly managed populations. Our results suggest management efforts can have substantial impacts on progeny quantity, but progeny quality is likely influenced by more difficult to manage population or species-specific traits such as tree size, plant sex ratio, and pollinator behavior.     

Environmental stress maintains trioecy in nematode worms

Sex is determined by chromosomes in mammals but it can be influenced by the environment in many worms, crustaceans, and vertebrates. Despite this, there is little understanding of the relationship between ecology and the evolution of sexual systems. The nematode Auanema freiburgensis has a unique sex determination system in which individuals carrying one X chromosome develop into males while XX individuals develop into females in stress-free environments and self-fertile hermaphrodites in stressful environments. Theory predicts that trioecious populations with coexisting males, females, and hermaphrodites should be unstable intermediates in evolutionary transitions between mating systems. In this article, we study a mathematical model of reproductive evolution based on the unique life history and sex determination of A. freiburgensis. We develop the model in two scenarios, one where the relative production of hermaphrodites and females is entirely dependent on the environment and one based on empirical measurements of a population that displays incomplete, “leaky” environmental dependence. In the first scenario environmental conditions can push the population along an evolutionary continuum and result in the stable maintenance of multiple reproductive systems. The second “leaky” scenario results in the maintenance of three sexes for all environmental conditions. Theoretical investigations of reproductive system transitions have focused on the evolutionary costs and benefits of sex. Here, we show that the flexible sex determination system of A. freiburgensis may contribute to population-level resilience in the microscopic nematode's patchy, ephemeral natural habitat. Our results demonstrate that life history, ecology, and environment may play defining roles in the evolution of sexual systems.     

Anisogamy,sexual selection,and the evolution and maintenance of sex

Summary In the present paper we distinguish between two aspects of sexual reproduction. Genetic recombination is a universal features of the sexual process. It is a primitive condition found in simple, single-celled organisms, as well as in higher plants and animals. Its function is primarily to repair genetic damage and eliminate deleterious mutations. Recombination also produces new variation, however, and this can provide the basis for adaptive evolutionary change in spatially and temporally variable environments.The other feature usually associated with sexual reproduction, differentiated male and female roles, is a derived condition, largely restricted to complex, diploid, multicellular organisms. The evolution of anisogamous gametes (small, mobile male gametes containing only genetic material, and large, relatively immobile female gametes containing both genetic material and resources for the developing offspring) not only established the fundamental basis for maleness and femaleness, it also led to an asymmetry between the sexes in the allocation of resources to mating and offspring. Whereas females allocate their resources primarily to offspring, the existence of many male gametes for each female one results in sexual selection on males to allocate their resources to traits that enhance success in competition for fertilizations. A consequence of this reproductive competition, higher variance in male than female reproductive success, results in more intense selection on males.The greater response of males to both stabilizing and directional selection constitutes an evolutionary advantage of males that partially compensates for the cost of producing them. The increased fitness contributed by sexual selection on males will complement the advantages of genetic recombination for DNA repair and elimination of deleterious mutations in any outcrossing breeding system in which males contribute only genetic material to their offspring. Higher plants and animals tend to maintain sexual reproduction in part because of the enhanced fitness of offspring resulting from sexual selection at the level of individual organisms, and in part because of the superiority of sexual populations in competition with asexual clones.     

Comparison of gametophyte growth, sex determination and reproduction in three fern species from the tropics

The gametophytes of Athyrium boryanurn, Thelypteris subpubescens and Pteridium aquilinum (vars. aquilinum, esculentum and wightianum) were grown as isolates, pairs and groups, and they were found to differ in their life histories: A. boryanum is a selfer which is sensitive to competition, while T. subpubescens is an ineffective selfer which is sensitive to competition, and P. aquilinurn is generally ineffective at sexual reproduction. A relationship between the level of gametophyte reproduction and the sporophyte viability was detected. The interactions between neighbours (competition or antheridiogen effects) appeared to influence gametophyte growth and reproductive success. In general, hermaphroditic gametophytes were considerably larger than males in all species, and those hermaphrodites which formed sporophytes were often smaller than other hermaphrodites.     

Extending pedigree analysis for uncertain parentage and diverse breeding systems

Breeding programs aimed at conserving genetic diversity in populations of wildlife or rare domestic breeds rely on detailed pedigree analysis for selection of breeders that will minimize the loss of alleles, reduce the accumulation of inbreeding, and maintain gene diversity. Commonly, techniques use a matrix of kinship coefficients to derive measures of genetic variation, inbreeding, and the value of individuals as breeders. Although these techniques were first developed for use on known pedigrees of diploid individuals, the concepts and methods can be extended to apply to any entity that contains genes derived from definable sources (e.g., individual parents, social groups, colonies, gene banks) via a definable mechanism of heredity (e.g., sexual reproduction between separate sexes, hermaphroditic selfing, autozygous production of homozygous or haploid offspring, cloning). Individuals with partly unknown ancestry or multiple possible parents can also be incorporated into kinship calculations, based on probabilistic assignment of parental contributions. This paper presents the algorithms used in new PMx software to extend traditional pedigree analysis techniques used for complete pedigrees of sexually reproducing, diploid species to deal with missing information due to unknown or uncertain parentage, and other breeding systems such as clones, selfing hermaphrodites, and haploid offspring or autogamy.     

Modern Portfolio Theory and the prudent hermaphrodite

Summary

Employment of Markowitz's Modern Portfolio Theory, economic models designed to predict the effect of variance and covariance on optimal investment allocation, may explain a wide variety of anomalies in reproductive biology. Natural selection appears to favor genetic diversity among offspring to a greater extent than is predicted by current theory. Consideration of the possible increase in fitness by reducing the covariance among offspring may help explain a variety of phenomena from multiple mating to the evolution of recombination (i.e., overcoming “the cost of meiosis”). Modern Portfolio Theory also make novel predictions as to when hermaphrodites should prefer the male vs. female role, i.e., engage in egg- vs. sperm-trading. It predicts that the sexual role with the lower variance in reproductive success will be preferred in hermaphrodites. This contradicts Bateman's principle that the male role is usually preferred due to energetic considerations but is consistent with Gillespie's principle. The available data suggest that mating hermaphrodites are risk-averse; gamete-trading whether of eggs or sperm is a strategy to reduce risk. In addition to overall variance, the skew of the distribution can be used to predict the mating systems of hermaphrodites and thus clarify the factors that are responsible for observed patterns of sex allocation and sexual conflict. The reduction of covariance among offspring may also help resolve “Williams' paradox”; that the observed distribution of dieoecy vs. simultaneous hermaphroditism in the Animal Kingdom cannot be explained by the prevailing models of the evolution of hermaphroditism.     

Reproductive assurance drives transitions to self-fertilization in experimental <Emphasis Type="Italic">Caenorhabditis elegans</Emphasis>

Background

Evolutionary transitions from outcrossing between individuals to selfing are partly responsible for the great diversity of animal and plant reproduction systems. The hypothesis of `reproductive assurance’ suggests that transitions to selfing occur because selfers that are able to reproduce on their own ensure the persistence of populations in environments where mates or pollination agents are unavailable. Here we test this hypothesis by performing experimental evolution in Caenorhabditis elegans.

Results

We show that self-compatible hermaphrodites provide reproductive assurance to a male-female population facing a novel environment where outcrossing is limiting. Invasions of hermaphrodites in male-female populations, and subsequent experimental evolution in the novel environment, led to successful transitions to selfing and adaptation. Adaptation was not due to the loss of males during transitions, as shown by evolution experiments in exclusively hermaphroditic populations and in male-hermaphrodite populations. Instead, adaptation was due to the displacement of females by hermaphrodites. Genotyping of single-nucleotide polymorphisms further indicated that the observed evolution of selfing rates was not due to selection of standing genetic diversity. Finally, numerical modelling and evolution experiments in male-female populations demonstrate that the improvement of male fitness components may diminish the opportunity for reproductive assurance.

Conclusions

Our findings support the hypothesis that reproductive assurance can drive the transition from outcrossing to selfing, and further suggest that the success of transitions to selfing hinges on adaptation of obligate outcrossing populations to the environment where outcrossing was once a limiting factor.

     

The evolution of phally polymorphism in Bulinus truncatus (Gastropoda,Planorbidae): the cost of male function analysed through life-history traits and sex allocation

In the hermaphrodite freshwater snail Bulinus truncatus, two sexual morphs, euphallic (regular hermaphrodites) and aphallic individuals without a male copulatory organ, co-occur at various ratios in natural populations. Both aphallic and euphallic individuals can reproduce by selfing, but when outcrossing aphallic individuals can only play the female role. A comparison of life-history traits and sex allocation in these two forms provides the opportunity to investigate the evolution and maintenance of sexual polymorphisms. This study was performed to test whether a reallocation of resources from the lost male function to the female function occurs in aphallic snails at the level of both sex organs (sex allocation) and life-history traits. In a first experiment we compared life-history traits over a whole life-cycle under selfing between the two sexual morphs. In a second experiment, the sex organs were weighed to test for a difference in sex allocation between the two morphs. No difference in resource allocation to female function between the two morphs was observed in either experiment. This is in contrast to patterns frequently observed in sexually polymorphic plants, and in a previous study performed on aphally in the same snail species. We discuss the genetic and physiological hypotheses that could explain these results, and their consequences for the evolution and maintenance of phally polymorphism in B. truncatus.