Mating system contributes only slightly to female maintenance in gynodioecious Geranium maculatum (Geraniaceae)

Gynodioecy, the co-occurrence of female and hermaphroditic individuals within a population, is an important intermediate in the evolution of separate sexes. The first step, female maintenance, requires females to have higher seed fitness compared with hermaphrodites. A common mechanism thought to increase relative female fitness is inbreeding depression avoidance, the magnitude of which depends on hermaphroditic selfing rates and the strength of inbreeding depression. Less well studied is the effect of biparental inbreeding on female fitness. Biparental inbreeding can affect relative female fitness only if its consequence or frequency differs between sexes, which could occur if sex structure and genetic structure both occur within populations. To determine whether inbreeding avoidance and/or biparental inbreeding can account for female persistence in Geranium maculatum, we measured selfing and biparental inbreeding rates in four populations and the spatial genetic structure in six populations. Selfing rates of hermaphrodites were low and did not differ significantly from zero in any population, leading to females gaining at most a 1–14% increase in seed fitness from inbreeding avoidance. Additionally, although significant spatial genetic structure was found in all populations, biparental inbreeding rates were low and only differed between sexes in one population, thereby having little influence on female fitness. A review of the literature revealed few sexual differences in biparental inbreeding among other gynodioecious species. Our results show that mating system differences may not fully account for female maintenance in this species, suggesting other mechanisms may be involved.     

FREQUENCY-DEPENDENT FITNESS IN SILENE VULGARIS,A GYNODIOECIOUS PLANT

In gynodioecious plants the selective processes that determine the relative number of female and hermaphroditic individuals are often frequency dependent. Frequency-dependent fitness can occur in the two sexes through a variety of mechanisms, especially given pollen limitation and inbreeding depression when hermaphrodites are rare. Frequency dependence in several components of the fitness of female and hermaphroditic Silene vulgaris was tested in experiments in which the relative numbers of the two sexes was varied among 12 artificial populations. In females, the proportion of flowers that set fruit covaried positively among populations with the frequency of hermaphrodites in two separate experiments, whereas the number of flowers/plant covaried negatively in one case. In hermaphrodites, the number of seeds/fruit covaried positively with the frequency of hermaphrodites, whereas the fitness of hermaphrodites estimated through pollen transfer covaried negatively. The results are discussed as they relate to the selective maintenance of gynodioecy in S. vulgaris and in light of a recent model of the effect of population structure on selection in gynodioecious systems.     

Floral dimorphism, pollination, and self-fertilization in gynodioecious GERANIUM RICHARDSONII (Geraniaceae)

The selective maintenance of gynodioecy depends on the relative fitness of the male-sterile (female) and hermaphroditic morphs. Females may compensate for their loss of male fitness by reallocating resources from male function (pollen production and pollinator attraction) to female function (seeds and fruits), thus increasing seed production. Females may also benefit from their inability to self-fertilize if selfing and inbreeding depression reduce seed quality in hermaphrodites. We investigated how differences in floral resource allocation (flower size) between female and hermaphroditic plants affect two measures of female reproductive success, pollinator visitation and pollen receipt, in gynodioecious populations of Geranium richardsonii in Colorado. Using emasculation treatments in natural populations, we further examined whether selfing by autogamy and geitonogamy comprises a significant proportion of pollen receipt by hermaphrodites. Flowers of female plants are significantly smaller than those of hermaphrodites. The reduction in allocation to pollinator-attracting structures (petals) is correlated with a significant reduction in pollinator visitation to female flowers in artificial arrays. The reduction in attractiveness is further manifested in significantly less pollen being deposited on the stigmas of female flowers in natural populations. Autogamy is rare in these protandrous flowers, and geitonogamy accounts for most of the difference in pollen receipt between hermaphrodites and females. Female success at receiving pollen was negatively frequency dependent on the relative frequency of females in populations. Thus, two of the prerequisites for the maintenance of females in gynodioecious populations, differences in resource allocation between floral morphs and high selfing rates in hermaphrodites, occur in G. richardsonii.     

Consequences of inbreeding for offspring fitness and gender in Silene vulgaris,a gynodioecious plant

Abstract In gynodioecious plants, hermaphrodite and female plants co‐occur in the same population. In these systems gender typically depends on whether a maternally inherited cytoplasmic male sterility factor (CMS) is counteracted by nuclear restorer alleles. These restorer alleles are often genetically dominant. Although plants of the female morph are obligatorily outcrossing, hermaphrodites may self. This selfing increases homozygosity and may thus have two effects: (1) it may decrease fitness (i.e. result in inbreeding depression) and (ii) it may increase homozygosity of the nuclear restorer alleles and therefore increase the production of females. This, in turn, enhances outcrossing in the following generation. In order to test the latter hypothesis, experimental crosses were conducted using individuals derived from four natural populations of Silene vulgaris, a gynodioecious plant. Treatments included self‐fertilization of hermaphrodites, outcrossing of hermaphrodites and females using pollen derived from the same source population as the pollen recipients, and outcrossing hermaphrodites and females using pollen derived from different source populations. Offspring were scored for seed germination, survivorship to flowering and gender. The products of self‐fertilization had reduced survivorship at both life stages when compared with the offspring of outcrossed hermaphrodites or females. In one population the fitness of offspring produced by within‐population outcrossing of females was significantly less than the fitness of offspring produced by crossing females with hermaphrodites from other populations. Self‐fertilization of hermaphrodites produced a smaller proportion of hermaphroditic offspring than did outcrossing hermaphrodites. Outcrossing females within populations produced a smaller proportion of hermaphrodite offspring than did crossing females with hermaphrodites from other populations. These results are consistent with a cytonuclear system of sex determination with dominant nuclear restorers, and are discussed with regard to how the mating system and the genetics of sex determination interact to influence the evolution of inbreeding depression.     

Maternal sex and mate relatedness affect offspring quality in the gynodioecious Silene acaulis

In gynodioecious species, females sacrifice fitness by not producing pollen, and hence must have a fitness advantage over hermaphrodites. Because females are obligately outcrossed, they may derive a fitness advantage by avoiding selfing and inbreeding depression. However, both sexes are capable of biparental inbreeding, and there are currently few estimates of the independent effects of maternal sex and multiple levels of inbreeding on female advantage. To test these hypotheses, females and hermaphrodites from six Alaskan populations of Silene acaulis were crossed with pollen from self (hermaphrodites only), a sibling, a random plant within the same population, and a plant from a different population. Germination, survivorship and early growth revealed inbreeding depression for selfs and higher germination but reduced growth in sib-crosses, relative to outcrosses. Independent of mate relatedness, females germinated more seeds that grew faster than offspring from hermaphrodites. This indicates that inbreeding depression as well as maternal sex can influence breeding system evolution. The effect of maternal sex may be explained by higher performance of female genotypes and a greater abundance of female genotypes among the offspring of female mothers.     

The Evolution of Dicliny in Schiedea (Caryophyllaceae), an Endemic Hawaiian Genus

Abstract The evolution of dioecy was studied in Schiedea (Caryophyllaceae), a genus endemic to the Hawaiian Islands. Eight of the 22 species are diclinous, possessing gynodioecious, subdioecious, or dioecious breeding systems. A biogeographic analysis of the genus indicates that the ancestor of Schiedea colonized early in the history of the Hawaiian Islands. Subsequently, hermaphroditic species appear to have engaged in inter-island colonization more frequently than diclinous species. For this reason, single-island endemism and dicliny are more common on the older Hawaiian Islands. Strong inbreeding depression was detected in three species of Schiedea , indicating that genetic factors have played a role in the evolution of dicliny. Depending on the level of natural selfing, the expression of inbreeding depressioin may have favored the outcrossed progeny of rare females in populations, and eventually the evolution of dioecy. In contrast to evidence for inbreeding depression, there was very little evidence that resource allocation, sex lability, or habitat partitioning have played an important role in the evolution of dioecy. In subdioecious S. globosa hermaphrodites were largely male in function, and in gynodioecious S. salicaria females and hermaphrodites were equivalent in nearly all aspects of female function that could be measured. Variation in breeding systems in Schiedea and the closely related Alsinidendron may result from the past history of population bottlenecks that have resulted in varying levels of inbreeding depression.     

INBREEDING DEPRESSION IN GYNODIOECIOUS LOBELIA SIPHILITICA: AMONG-FAMILY DIFFERENCES OVERRIDE BETWEEN-MORPH DIFFERENCES

If inbreeding depression is caused by deleterious recessive alleles, as suggested by the partial dominance hypothesis, a negative correlation between inbreeding and inbreeding depression is predicted. This hypothesis has been tested several times by comparisons of closely related species or comparisons of populations of the same species with different histories of inbreeding. However, if one is interested in whether this relationship contributes to mating-system evolution, which occurs within populations, comparisons among families within a population are needed; that is, inbreeding depression among individuals with genetically based differences in their rate of selfing should be compared. In gynodioecious species with self-compatible hermaphrodites, hermaphrodites will have a greater history of potential inbreeding via both selfing and biparental inbreeding as compared to females and may therefore express a lower level of inbreeding depression. We estimated the inbreeding depression of female and hermaphrodite lineages in gynodioecious Lobelia siphilitica in a greenhouse experiment by comparing the performance of selfed and outcrossed progeny, as well as sibling crosses and crosses among subpopulations. We did not find support for lower inbreeding depression in hermaphrodite lineages. Multiplicative inbreeding depression (based on seed germination, juvenile survival, survival to flowering, and flower production in the first growing season) was not significantly different between hermaphrodite lineages (δ = 0.30 ± 0.08) and female lineages (δ = 0.15 ± 0.18), although the trend was for higher inbreeding depression in the hermaphrodite lineages. The population-level estimate of inbreeding depression was relatively low for a gynodioecious species (δ = 0.25) and there was no significant inbreeding depression following biparental inbreeding (δ = 0.01). All measured traits showed significant variation among families, and there was a significant interaction between family and pollination treatment for four traits (germination date, date of first flowering, number of flowers, and aboveground biomass). Our results suggest that the families responded differently to selfing and outcrossing: Some families exhibited lower fitness following selfing whereas others seemed to benefit from selfing as compared to outcrossing. Our results support recent simulation results in that prior inbreeding of the lineages did not determine the level of inbreeding depression. These results also emphasize the importance of determining family-level estimates of inbreeding depression, relative to population-level estimates, for studies of mating-system evolution.     

Breeding systems of hermaphroditic and gynodioecious populations of the colonizing species Trifolium hirtum All. in California

Summary A multilocus procedure was used to estimate outcrossing rates in ten roadside populations of Trifolium hirtum in California. Three groups of populations were studied: cultivars, hermaphroditic, and gynodioecious (sexually dimorphic) populations. The multilocus outcrossing rate (t_m) varied from 0.05 to 0.43 among populations. Population level t_m estimates were significantly correlated with the observed heterozygosity in gynodioecious populations but not in hermaphroditic populations. The outcrossing rate of hermaphrodites and females was estimated in three gynodioecious populations; the estimates of t_m varied from 0.09 to 0.23 for hermaphrodites and from 0.73 to 0.80 for females. The distribution of outcrossing rates in gynodioecious populations is bimodal. Our results indicate that for the levels of selfing observed among hermaphrodites, inbreeding depression is likely to be a major factor in the maintenance of females in gynodioecious populations.     

FEMALE FREQUENCIES IN GYNODIOECIOUS POPULATIONS CORRELATED WITH SELFING RATES IN HERMAPHRODITES

Gynodioecious populations consist of separate hermaphroditic and female individuals. Females are at a selective disadvantage because they contribute genes to the next generation only through ovules, while hermaphrodites contribute genes through ovules and pollen. For females to be maintained in populations they must have some compensating selective advantage. The outcrossing hypothesis postulates that females are maintained because their progeny result from obligate outcrossing, whereas some of the progeny of hermaphrodites result from self-fertilization and are less fit because of inbreeding depression. If correct, the frequency of females should be positively correlated with selfing rates of hermaphrodites in populations. We found a strong positive correlation between female frequency and selfing rates of hermaphrodites (r = 0.91, P < 0.01) in eight gynodioecious populations of Hawaiian species of Bidens. Our results confirm that the obligate outcrossing of females is a major factor maintaining females in gynodioecious populations. However, the observed selfing rates are insufficient by themselves to account for the frequency of females in these populations.     

Sex allocation of females and hermaphrodites in the gynodioecious Geranium sylvaticum

Seed production and patterns of sex allocation were studied in female and hermaphroditic plants in two gynodioecious populations of Geranium sylvaticum (Geraniaceae). Females produced more flower buds and seeds than hermaphrodites in one of the two study populations. The other female traits measured (pistil biomass, seed number per fruit, individual seed mass) did not differ between the gender morphs. The relative seed fitness of hermaphrodites differed between the study populations, with hermaphrodites gaining less of their fitness through female function in the population with a high frequency of females. However, the amount and size of pollen produced by hermaphrodites did not differ between populations. The number of flower buds was positively correlated with seed production in females, whereas in hermaphrodites a positive correlation between number of buds and seed production was found in only one of the two study populations. These results suggest that fitness gain through female function is labile in hermaphrodites of this species, and is probably affected by environmental factors such as the sex ratio of the population.     

HOW SELFING,INBREEDING DEPRESSION,AND POLLEN LIMITATION IMPACT NUCLEAR‐CYTOPLASMIC GYNODIOECY: A MODEL

Gynodioecy, the co‐occurrence of females and hermaphrodites, is often due to conflicting interactions between cytoplasmic male sterility genes and nuclear restorers. Although gynodioecy often occurs in self‐compatible species, the effect of self‐pollination, inbreeding depression, and pollen limitation acting differently on females and hermaphrodites remains poorly known in the case of nuclear‐cytoplasmic gynodioecy (NCG). In this study, we model NCG in an infinite population and we study the effect of selfing rate, inbreeding depression, and pollen limitation on the maintenance of gynodioecy and on sex ratios at equilibrium. We found that selfing and inbreeding depression have a strong impact, which depends on whether restorer cost acts on male or female fitness. When cost affects male fitness, the strength of cost has no effect, whereas selfing and inbreeding depression only impact gynodioecy by modifying the value of the female advantage. When cost affects female fitness, selfing facilitates NCG and reduces the role of strength of the cost, even when no inbreeding depression occurs, whereas inbreeding depression globally restricts the maintenance of the polymorphism. Finally, we found that pollen limitation could additionally strongly modify the dynamic of gynodioecy. We discuss our findings in the light of empirical data available in gynodioecious species.     

AN ECOLOGICAL GENETIC STUDY OF GYNODIOECY IN LIMNANTHES DOUGLASII (LIMNANTHACEAE)

Gynodioecy in two populations of Limnanthes douglasii var. rosea was studied for its maintenance requirements, role in population structure and its influence on the levels of outbreeding. The mode of inheritance of male sterility appeared to be nucleo-cytoplasmic, although the precise number of nuclear genes and dominance relationships could not be ascertained. Measurements on the plants sampled from natural stands and controlled experiments showed that male-steriles and their progeny had greater biomass and more flowers per plant than hermaphrodites, though these results varied with the environmental conditions. The hermaphrodites of gynodioecious populations had higher rates of selfing than the hermaphrodite individuals in populations lacking male sterility. Estimates of the inbreeding depression and homozygosity levels were also higher in gynodioecious populations. Variation in these parameters of breeding system and relative heterozygosis among populations may explain why male sterility has a restricted distribution, as theoretical models also predict rather specific conditions for this stable polymorphism. These data suggest that the advantage of male sterility is associated with lowered inbreeding depression. However, the potential ecological resource reallocation to the female function needs to be investigated. The fitness differences observed in this study between male-steriles and hermaphrodites appear inadequate to maintain the nucleo-cytoplasmic male sterility but could account for the observed frequencies (10–20%) of male-steriles in nature if the genetic system has evolved from a cytoplasmic system.     

Testing for sex differences in biparental inbreeding and its consequences in a gynodioecious species

In many gynodioecious species cross-pollinated seeds from females outperform those from hermaphrodites. Using the gynodioecious alpine perennial Silene acaulis, I investigated whether this was the result of greater biparental inbreeding among hermaphrodites leading to greater biparental inbreeding depression. I also determined the influence of relatedness on progeny fitness. Experiments were performed using individuals from a site whose population structure and coefficient of inbreeding was known. In the first experiment, crosses were made on plants in the field to determine the effect of seven different crossing distances, plus selfing, on germination and early seedling survival and growth. Although selfed seeds died more often and grew slower than crossed seeds, the effect of crossing distance was negligible for all measured fitness traits, refuting the biparental inbreeding hypothesis as a mechanism to explain why seeds from hermaphrodites die more often than those from females. Nonetheless, cross-pollinated seeds from hermaphrodites did die more, indicating that another mechanism must be responsible. In the second experiment, the effect of different levels of inbreeding on germination and seedling survival was determined by growing seeds from experimental matings varying in relatedness. Inbreeding depression for a multiplicative fitness estimate was significant for all levels of inbreeding, suggesting that inbred individuals are unlikely to become established in the population and providing insight into the results of the first experiment. Alternative hypotheses are discussed to explain why seeds from hermaphrodites die more often, which together with the results of this study, suggest that the restoration of male function in hermaphrodites comes with a correlated cost to seedling survival.     

Inbreeding effects on progeny sex ratio and gender variation in the gynodioecious Silene vulgaris (Caryophyllaceae)

In gynodioecious species, sex expression is generally determined through cytoplasmic male sterility genes interacting with nuclear restorers of the male function. With dominant restorers, there may be an excess of females in the progeny of self-fertilized compared with cross-fertilized hermaphrodites. Moreover, the effect of inbreeding on late stages of the life cycle remains poorly explored. Here, we used hermaphrodites of the gynodioecious Silene vulgaris originating from three populations located in different valleys in the Alps to investigate the effects of two generations of self- and cross-fertilization on sex ratio and gender variation. We detected an increase in females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for female and male fertility. Male fertility correlated positively with sex ratio differences between outbred and inbred progeny, suggesting that dominant restorers are likely to influence male fertility qualitatively and quantitatively in S. vulgaris. We argue that the excess of females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for gamete production may contribute to the maintenance of females in gynodioecious populations of S. vulgaris because purging of the genetic load is less likely to occur.     

ECOLOGICAL GENETICS OF GYNODIOECY IN SILENE VULGARIS: RELATIVE FITNESS OF FEMALES AND HERMAPHRODITES DURING THE COLONIZATION PROCESS

Recent theoretical models have addressed the influence of metapopulation dynamics on the fitness of females and hermaphrodites in gynodioecious plants. In particular, selection is thought to favor hermaphrodites during population establishment because that sex should be less prone to pollen limitation, especially if self-fertilization is possible. However, inbreeding depression could limit this advantage. In this experimental study of Silene vulgaris, a weedy gynodioecious plant, the fitness of females and hermaphrodites was estimated from seed production in both mixed-sex populations and for individuals isolated from these populations by 20, 40, 80, or 160 m. In mixed populations females display statistically significant greater per capita seed production owing to higher capsule production and higher rates of seed germination. The fitness of both sexes declines with increasing isolation, but at different rates, such that in the 160-m treatment hermaphrodites are by far the more fit sex. Allozyme studies suggest that this differential decline is because the selfing rate in hermaphrodites increases as a function of isolation, at least partially compensating for a decline in the availability of outcross pollen. Overall, the negative effects of pollen limitation on females far outweighs the negative effects of inbreeding depression following selfing in hermaphrodites. Thus, extinction/recolonization dynamics would appear to favor hermaphrodites as long as seed dispersal events exceed some critical distance.     

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.     

The transition to gender dimorphism on an evolutionary background of self-incompatibility: an example from Lycium (Solanaceae)

Populations of three North American species of Lycium (Solanaceae) are morphologically gynodioecious and consist of male-sterile (i.e., female) and hermaphroditic plants. Marked individuals were consistent in sexual expression across years and male sterility was present throughout much of the species' ranges. Controlled pollinations reveal that L. californicum, L. exsertum, and L. fremontii are functionally dioecious. Fruit production in females ranged from 36 to 63%, whereas hermaphrodites functioned essentially as males. Though hermaphrodites were mostly male, investigation of pollen tube growth reveals that hermaphrodites of all dimorphic species were self-compatible. Self-fertilization and consequent inbreeding depression are commonly invoked as important selective forces promoting the invasion of male-sterile mutants into cosexual populations. A corollary prediction of these models is that gender dimorphism evolves from self-compatible ancestors. However, fruit production, seed production, and pollen tube number following outcross pollination were significantly higher than following self-pollination for three diploid, cosexual species that are closely related to the dimorphic species. The data presented here on incompatibility systems are consistent with the hypothesis that polyploidy disrupted the self-incompatibility system in the gynodioecious species leading to the evolution of gender dimorphism.     

Selfing and resource allocation in Schiedea salicaria (Caryophyllaceae), a gynodioecious species

Abstract Levels of selfing and resource allocation patterns were investigated in Schiedea salicaria (Caryophyllaceae), a gynodioecious species with high levels of inbreeding depression and nuclear control of male sterility. Selfing levels were higher in hermaphrodites than females, especially when adjusted for early acting inbreeding depression. The sexes of S. salicaria were similar in most allocation patterns including number of flowers and capsules per inflorescence, seeds per flower, and seed mass. Seeds produced by females had higher levels of germination than seeds of hermaphrodites, a likely result of high selfing levels and the expression of inbreeding depression in the progeny of hermaphrodites. Invasion of females in populations of S. salicaria is probably related to the expression of inbreeding depression at germination and in later life history stages. Comparisons with related species of Schiedea that also have nuclear control of male sterility suggest that reallocation of resources in hermaphrodites to male function occurs as females increase in frequency, but that resource reallocation is not important for the success of females when they first invade populations.     

Exploring the genetic basis and proximate causes of female fertility advantage in gynodioecious Thymus vulgaris

In many gynodioecous species, females produce more viable seeds than hermaphrodites. Knowledge of the relative contribution of inbreeding depression in hermaphrodites and maternal sex effects to the female fertility advantage and the genetic basis of variation in female fertility advantage is central to our understanding of the evolution of gender specialization. In this study we examine the relative contribution of inbreeding and maternal sex to the female fertility advantage in gynodioecious Thymus vulgaris and quantify whether there is genetically based variation in female fertility advantage for plants from four populations. Following controlled self and outcross (sib, within-population, and between-population) pollination, females had a more than twofold fertility advantage (based on the number of germinating seeds per fruit), regardless of the population of origin and the type of pollination. Inbreeding depression on viable seed production by hermaphrodites occurred in two populations, where inbreeding had been previously detected. Biparental inbreeding depression on viable seed production occurred in three of four populations for females, but in only one population for hermaphrodites. Whereas the maternal sex effect may consistently enhance female fertility advantage, inbreeding effects may be limited to particular population contexts where inbreeding may occur. A significant family x maternal sex interaction effect on viable seed production was observed, illustrating that the extent of female fertility advantage varies significantly among families. This result is due to greater variation in hermaphrodite (relative to female) seed fertility between families. Despite this genetic variation in female fertility advantage and the highly female biased sex ratios in populations of T. vulgaris, gynodioecy is a stable polymorphism, suggesting that strong genetic and/or ecological constraints influence the stability of this polymorphism.     

Do inbreeding depression and relative male fitness explain the maintenance of androdioecy in white mangrove, Laguncularia racemosa (Combretaceae)?

Mathematical models predict that to maintain androdioecious populations, males must have at least twice the fitness of male function in hermaphrodites. To understand how androdioecy is maintained in Laguncularia racemosa (white mangrove), outcrossing, inbreeding depression, and relative male fitness were estimated in two androdioecious populations and one hermaphroditic population. Outcrossing was estimated based on length of pollinator foraging bout and pollen carryover assumptions. Inbreeding depression was measured at three life stages: fruit set, seedling emergence, and seedling survivorship. The relative fitnesses of males and the male component of hermaphrodites were compared at these three stages and at the pollen production stage. Male frequency predictions generated by Lloyd's model were compared with observed frequencies in two androdioecious subpopulations. Outcrossing estimates were moderate for all populations (0.29-0.66). Inbreeding depression varied among populations (-0.03-0.86), but the strength of inbreeding depression did not increase with male frequency. Males produced significantly more flowers/inflorescence than hermaphrodites, but pollen production/flower did not differ. Male and hermaphroditic progeny did not differ significantly at other life stages. Populations of white mangrove with male plants were functionally androdioecious. Lloyd's model accurately predicted male frequency in one androdioecious subpopulation, but underestimated male frequency in the second subpopulation.