Mixed Mating Strategy Research Articles

Ruellia blechum: A new case of cleistogamy in Acanthaceae

AbstractCleistogamy is characterized by the presence of flowers that are permanently closed, yet they still produce fruits and seeds through autonomous self‐pollination. Populations with cleistogamous flowers can also have flowers that open, called chasmogamous, which promote cross‐pollination and genetic variability. Acanthaceae is among the families with the highest frequency of cleistogamy, observed mainly in Ruellia. This study aimed to assess cleistogamy in Ruellia blechum for the first time. In a population in Central Brazil, we analysed the production of cleistogamous and chasmogamous flowers, their fruiting, differences in flower bud size, self‐pollination in cleistogamous flowers and visitors in chasmogamous flowers. Of the flower buds observed, 35.29% remained closed (cleistogamous flowers), with 31.25% of them forming fruit. In contrast, 60.24% of the flowers that opened (chasmogamous) developed fruit. The length of cleistogamous flowers was shorter than that of chasmogamous floral buds in pre‐anthesis. On average, 24.45 pollen grains were self‐deposited on the stigma of a cleistogamous flower. The chasmogamous flowers were visited mainly by small bees. This study is the first to confirm the presence of cleistogamous flowers in R. blechum. However, their co‐occurrence with chasmogamous flowers suggests a fitness advantage for maintaining a mixed mating strategy.

Competition and drought affect cleistogamy in a non-additive way in the annual ruderal Lamium amplexicaule.

Competition affects mixed-mating strategies by limiting available abiotic or biotic resources such as nutrients, water, space, or pollinators. Cleistogamous species produce closed (cleistogamous, CL), obligately selfed, simultaneously with open (chasmogamous, CH), potentially outcrossed flowers. The effects of intraspecific competition on fitness and cleistogamy variation can range from limiting the production of costly CH flowers because of resource limitation, to favouring CH production because of fitness advantages of outcrossed, CH offspring. Moreover, the effects of competition can be altered when it co-occurs with other environmental variations. We grew plants from seven populations of the ruderal Lamium amplexicaule, originating from different climates and habitats, in a common garden experiment combining drought, interspecific competition, and seasonal variation. All these parameters have been shown to influence the degree of cleistogamy in the species on their own. In spring, competition and drought negatively impacted fitness, but the CL proportion only increased when plants were exposed to both treatments combined. We did not observe the same results in autumn, which can be due to non-adaptive phenotypic variation, or to differences in soil compactness between seasons. The observed responses are largely due to phenotypic plasticity, but we also observed phenotypic differentiation between populations for morphological, phenological, and cleistogamy traits, pointing to the existence of different ecotypes. Our data do not support the hypothesis that CL proportion should decrease when resources are scarce, as plants with reduced growth had relatively low CL proportions. We propose that variation in cleistogamy could be an adaptation to pollinator abundance, or to environment-dependent fitness differences between offspring of selfed and outcrossed seeds, two hypotheses worth further investigation. This opens exciting new possibilities for the study of the maintenance of mixed-mating systems using cleistogamous species as models that combine the effects of inbreeding and reproductive costs.

Contradistinctive floral attributes, pollination guilds and their consequence on the outcrossing rate in two elevational morphs of Rhododendron arboreum Sm.

Infraspecific floral trait variations may appear in response to elevational differences in alpine plant species. There is enormous information on the selection of such morphs mediated by biotic and/or abiotic variables. Whether such differences contribute to differences in reproductive strategy and mating outcomes is rarely investigated. We investigated these aspects in two distinct elevational floral morphs (Red and Pink) of Rhododendron arboreum Sm. in Western Himalaya. The red morphs occupy the lower elevations while pink morphs the higher elevations. The two morphs differ in floral traits like phenology, dimension, display, quality of floral rewards, and pollinators that happen to influence interaction with available pollinator pool at each elevation. The pink morph exhibits entomophily, while the red ones show ornithophily. Although experimental pollinations established that both the morphs are self-compatible, selfing results in significantly lower fruit-set than either cross- or open-pollinations. The outcrossing rate in the red morph, as determined by using simple sequence repeat (SSR) markers, was higher (tm=0.82) than that in the pink morph (tm=0.76), with a tendency of the latter to be shifting towards mixed-mating strategy. However, the extent of biparental inbreeding was comparable among the two morphs. It is inferred that the differences in the mating outcomes among the morphs in the tree species are linked to those emerging from floral traits and the pollination by different functional groups of floral visitors.

Epigenetic mechanisms of DNA methylation in the transgenerational effect of ethylhexyl salicylate on zebrafish

In this study, a 120-day whole-life cycle exposure and oviposition experiment on zebrafish with maternal and paternal mixed mating strategy was conducted to investigate the epigenetic mechanism of DNA methylation in ethylhexyl salicylate (EHS, 1, 10, 100 μg/L)-induced transgenerational effects. Results showed that EHS could induce the decrease of DNA methyltransferase 1 (DNMT1) activity and average global DNA methylation level in maternal parents and the increase of the above indexes in paternal parents, while the change of glycine N-methyltransferase activity was opposite to DNMT1. The average global DNA methylation levels were significantly increased in the offsprings of both parents exposed and father-only exposed to EHS, suggesting that EHS-induced epigenetic modifications may be stable and heritable. Hierarchical clustering analysis of promoter at different methylation sites showed that the DNA methylation pattern of offsprings were similar to that of the paternal parents, meaning that the offsprings may have inherited paternal DNA methylation pattern with eya2, pcdh2g5 and pcdh2g1 as key genes and lead to high locomotor activity in offsprings. KEGG pathway analysis showed that parental exposure to EHS may interfere with the central nervous system, insulin function system, melanogenesis system and the normal development of somatic axis of offsprings.

Mixed pollination system and floral signals of Paepalanthus (Eriocaulaceae): insects and geitonogamy ensure high reproductive success.

Eriocaulaceae exhibit a great variety of floral traits associated with insect (e.g. nectariferous structures) and wind pollination (unisexual flowers, exposed sexual organs and small pollen grains), as well as the 'selfing syndrome' (small flowers, short distance between stigma and anthers, and temporal overlap of male and female phases). Paepalanthus bifidus, P. subtilis and P. tortilis are related species that differ in form, size and colour of floral structures. We aimed to investigate the pollination and reproductive biology of these three species. We analysed the floral biology, floral visitors, pollinator behaviour, and the contribution of insects, wind and spontaneous geitonogamy to fruit set. We also evaluated the floral colour and scent of the species. Colour reflectance of capitula of each species was measured and plotted in models of insect vision. Floral scent samples were extracted and the compounds were compared to vegetative scent samples. In all species, the staminate and pistillate flowers are arranged in alternating cycles with a temporal overlap between these phases. Ants were the most frequent floral visitors and were effective pollinators in P. bifidus and P. tortilis, while flies were occasional pollinators in P. tortilis. Floral visitors were not observed in P. subtilis. In all species, fruits were produced by spontaneous geitonogamy, with no evidence of wind pollination. According to the models of insect vision, the colours of the capitula of P. bifidus and P. subtilis are the most inconspicuous for ants and flies. We found no difference between the emission of volatiles of inflorescences and vegetative structures. This study suggests that ant pollination might be more widespread in Eriocaulaceae than currently assumed. Furthermore, for small monocarpic plants, mixed mating strategies are most favourable, by ensuring reproduction either by outcrossing when pollinators are abundant or by spontaneous geitonogamy when pollinations are scarce/absent.

Fitness dependence preserves selection for recombination across diverse mixed mating strategies

Meiotic recombination and the factors affecting its rate and fate in nature have inspired many studies in theoretical evolutionary biology. Classical theoretical models have inferred that recombination can be favored under a rather restricted parameter range. Thus, the ubiquity of recombination in nature remains an open question. However, these models assumed constant recombination with an equal rate across all individuals within the population, whereas empirical evidence suggests that recombination may display certain sensitivity to ecological stressors and/or genotype fitness. Models assuming condition-dependent recombination show that such a strategy can often be favored over constant recombination. Moreover, in our recent model with panmictic populations subjected to purifying selection, fitness-dependent recombination was quite often favored even when any constant recombination was rejected. By using numerical modeling, we test whether such a ‘recombination-rescuing potential’ of fitness dependence holds also beyond panmixia, given the recognized effect of mating strategy on the evolution of recombination. We show that deviations from panmixia generally increase the recombination-rescuing potential of fitness dependence, with the strongest effect under intermediate selfing or high clonality. We find that under partial clonality, the evolutionary advantage of fitness-dependent recombination is determined mostly by selection against heterozygotes and additive-by-additive epistasis, while under partial selfing, additive-by-dominance epistasis is also a driver.

Distribution Characteristics, Population Structure and Dynamics of the Endangered Plant, <i>Viola websteri</i> Hemsl.

Plant species exhibit current characteristics as a result of interactions with environmental conditions. The plants of Viola sp. have selected chasmogamous flowers with vigorous vegetative propagation or development of cleistogamous flowers as an adaptation strategy. Viola websteri is distributed on the Korean peninsula and the eastern part of Jilin Province, China. The center and edge of the distribution are expected to exhibit different population-dynamics. It is necessary to investigate the cause of its current limited distribution even though V. websteri has a mixed-mating strategy. Firstly, We examined the vegetation environment of habitats and evaluated its characteristics. Growth characteristics were examined through plant phenology. We then evaluated the population structure, characteristics of chasmogamous flowers, and productivity of cleistogamous flowers. Moreover, we compared population sizes between 2014 and 2018. Most habitats were located in deciduous broadleaf mixed forests adjacent to valleys. V. websteri produced chasmogamous flowers with self-incompatibility in April-May and cleistogamous flowers in June-September. The cleistogamous flower production is a strategy ensuring seed production under uncertain environmental fluctuations; these were approximately twice as numerous as chasmogamous flowers. The population structure was distinguished into stable and very unstable regions. There were sites where the population experienced a sharp decline in the 2018 compared to that of 2014. This large decline was found in the edge populations. The habitats had different microsites depending on the natural disturbances of drought and the matrix constituting the habitat, thus supporting various plants. Ensuring the production of seeds through cleistogamous flowers, it was determined that rapid seedling re-establishment and population replenishment were possible when the natural disturbance factor was removed. Environmental factors did not equally affect all populations or individuals. Therefore, it was expected that it would be able to persisted in a long time, despite the rapid decrease in the number of individuals in the population regionally. Local extinction and re-establishment are likely to repeat according to environmental change. We propose the additional population investigation based on this works are required. We also suggest a need to assess the long-term population dynamics and the genetic characteristics of chasmogamous flowers and cleistogamous flowers to establish and implement effective conservation strategies.

Absence of strict monogamy in the Eurasian jackdaw, Coloeus monedula

Abstract The Eurasian Jackdaw is thought to be archetypically monogamous, but recent tagging research uncovered extra-pair copulations in the species. Here we examined extra-pair paternity (genetic monogamy) in Eurasian jackdaws breeding in the Judean Hills, Israel, at the global edge of the species range, using a set of highly polymorphic molecular microsatellites. We found roughly a sixth of nests sampled showed deviations from monogamy, more than previously found in DNA fingerprinting studies of jackdaws, suggesting a mixed mating strategy in this population. These findings support the trend of extra-pair paternity in avian species, even when social monogamy remains the rule, and highlight the importance of continued study of species throughout their geographical range.

Plurality in mating: Exploring the occurrence and contingencies of mating strategies

People adopt a variety of strategies in order to achieve specific mating goals. The current research nominated nine different mating strategies, and attempted to estimate their occurrence. Evidence from an online sample of 6273 Greek-speaking participants, indicated that a mixed strategy was in the highest occurrence, followed by a long-term and a short-term mating strategy. Men were more likely than women to prefer a short-term and a mixed mating strategy, and that younger participants were more likely to prefer a mixed than a long-term mating strategy. In addition, heterosexual women with same-sex attraction were more likely than exclusively heterosexual women to prefer a short-term and a mixed strategy than a long-term mating strategy. Furthermore, we found that men were more likely than women and older participants were more likely than younger participants to indicate that they would cheat on their partners if they were in a long-term intimate relationship. Furthermore, heterosexual with same-sex attraction, bisexual and homosexual men and women were more likely than exclusive heterosexual participants to indicate that they would cheat on their partners when in a long-term intimate relationship.

Mating Strategies: Exploring Sex Differences and Similarities in the Greek Cultural Context

People employ at least three strategies when they engage in mating: They look for a long-term partner (long-term mating strategy), short-term partners (short-term mating strategy), or a long-term partner along with casual extra-pair ones (mixed mating strategy). The current research examines the degree to which the two sexes diverge in the adoption of these strategies. In an online sample of 2,307 Greek-speaking participants, it was found that men were more likely than women to prefer a short-term strategy or a mixed strategy rather than a long-term strategy. However, the two sexes were similar in that large majorities of both preferred a long-term strategy. It was also found that a short-term strategy was preferred more frequently by younger than by older participants, and a mixed strategy was reported more frequently by married participants than by those who were in a relationship or who were single.

Clonal diversity and spatial genetic structure in the long-lived herb, Prairie trillium.

Combining population genetic studies with demographic surveys in long-lived clonal herbs can yield insight into the population dynamics of clonal plant populations. In this study, we assayed clonal diversity and spatial genetic structure in a population of a long-lived understory herb, Trillium recurvatum, that has been the focus of a demographic study spanning 26 years at the Meeman Biological Station in Memphis, Tennessee, USA. Using a set of five newly developed simple sequence repeat markers first reported here, we assessed 1) the extent of clonal diversity within the Meeman site, 2) the degree to which genetic diversity varies with stage class (juvenile, non-flowering, and flowering adults) at this site, 3) whether there is spatial genetic structure at the Meeman site, and 4) how measures of genetic diversity and inbreeding at the Meeman site compare to two additional nearby populations. Along with these analyses, we calculated and compared traditional population genetic metrics with information theory-based diversity indices. Although clonal propagation was present, the focal population displayed moderate levels of clonal diversity comprising 81 genets from the 174 individuals sampled. In the focal site, we also found that genetic diversity was highest in the flowering stage class when compared to the non-flowering and juvenile classes. We report that genets exhibited spatial genetic structure in the focal site exhibiting values for the Sp statistic of 0.00199 for linear distance and 0.0271 for log distance. Measures of unbiased gene diversity and the inbreeding coefficient were comparable across the sampled populations. Our results provide complementary genetic data to previous demographic studies in T. recurvatum, and these findings provide data for future studies aimed at integrating the degree of clonality, genetic variation, and population dynamics in this species. Our findings suggest that T. recurvatum at the focal Meeman site displays higher levels of sexual reproduction than were previously suggested, and spatial genetic structure estimates were comparable to other plant species with mixed and outcrossing mating strategies.

Development of microsatellite markers for the annual andromonoecious herb Commelina communis f. ciliata (Commelinaceae).

Commelina communis f. ciliata (Commelinaceae), a newly distinguished taxon, is an annual andromonoecious herb exhibiting a mixed mating system, the details of which remain unclear. We developed microsatellite markers for use in exploring the evolution of andromonoecy and mixed mating in the species. Fifteen microsatellite loci were developed using next-generation sequencing. The primer sets were used to evaluate 65 C. communis f. ciliata individuals from three populations in Japan; we found 1-13 alleles per locus and the expected heterozygosity ranged from 0.00 to 0.76. The markers are potentially useful to examine intra- and interspecies genetic structure and the mixed mating strategy of Commelina species via paternity analysis.

Extra‐pair offspring are less heterozygous than within‐pair offspring in American redstarts Setophaga ruticilla

The vast majority of bird species are socially monogamous; however, extra‐pair paternity is nearly ubiquitous and a number of theories have been proposed to explain the prevalence of this mixed mating strategy. Here, we test the genetic compatibility hypothesis – the idea that females seek extra‐pair copulations with males whose genes are more compatible with her own. For this study, we examined eight years of paternity data (2004–2011) from a Nearctic‐Neotropical migratory bird, the American redstart Setophaga ruticilla, breeding in southeastern Ontario, Canada. We predicted that females paired with genetically similar males (higher relatedness) would be more likely to produce extra‐pair offspring and that extra‐pair offspring would have higher levels of heterozygosity than within‐pair offspring. Alternatively, because this population experiences high levels of immigration, females may produce extra‐pair offspring with more genetically similar males because of the potential for outbreeding depression. Using five highly variable microsatellite markers, we examined patterns of relatedness among social pairs as well as measures of offspring heterozygosity. In contrast to our predictions, we found no difference in relatedness between social pairs where the females produced extra‐pair offspring and social pairs where the females produced only within‐pair offspring. However, extra‐pair offspring were significantly less heterozygous than within‐pair offspring. Together, these findings suggest that females a) are not engaging in extra‐pair fertilizations based on relatedness to their social mate and b) appear to be mating with extra‐pair males that are more genetically similar to themselves. We suggest there may be benefits for females to mate with genetically similar extra‐pair males in highly outbred populations with high rates of immigration, such as for maintaining co‐adapted gene complexes or genes coding for local adaptations.

Pollination biology and breeding system in five nocturnal species of Oenothera (Onagraceae): reproductive assurance and opportunities for outcrossing

The capacity to produce seed, both by selfing and outcrossing, or mixed mating strategies, is considered a mechanism for overcoming unpredictable pollinator availability. In the present study, we investigate breeding system, insect visitations and the role of insect visitors in the pollination of five species of Oenothera subsect. Oenothera. Field experiments showed that autonomous selfing occurs at bud stage, prior to the opening of the flower. Control flowers showed similar seed set to hand-pollinated flowers, whereas emasculated flowers and those subject to open pollination set fewer seed. These results are consistent with the hypothesis that the investigated Oenothera exhibit a great capacity for autonomous selfing and that selfing is selected in order to provide reproductive assurance. Although flowers were visited mostly by nocturnal lepidopterans, these insects did not precipitate pollination and are thus considered nectar thieves. Conversely, analysis of pollen loads and behavior during foraging by diurnal insect visitors revealed that honeybees and bumblebees are the probable pollinators. We conclude that production of flowers capable of autonomous selfing at bud stage, followed by anthesis and opportunities for outcrossing, probably improves the invasive potential of these Oenothera in Europe, together with a rapid increase in their populations, even when pollinators are scarce.

Floral contrivances and specialised pollination mechanism strongly influence mixed mating in Wrightia tomentosa (Apocynaceae).

Reproductive success of a plant species is largely influenced by the outcome of mating pattern in a population. It is believed that a significantly larger proportion of animal-pollinated plants have evolved a mixed-mating strategy, the extent of which may vary among species. It is thus pertinent to investigate the key contributors to mating success, especially to identify the reproductive constraints in depauperate populations of threatened plant species. We examined the contribution of floral architecture, pollination mechanism and breeding system on the extent of outcrossing rate in a near-threatened tree species, Wrightia tomentosa. The breeding system was ascertained from controlled pollination experiments. In order to determine outcrossing rate, 60 open-pollinated progeny were analysed using an AFLP markers. Although the trees are self-compatible, herkogamy and compartmentalisation of pollen and nectar in different chambers of the floral tube effectively prevent spontaneous autogamy. Pollination is achieved through specialised interaction with moths. Differential foraging behaviour of settling moths and hawkmoths leads to different proportions of geitonogamous and xenogamous pollen on the stigma. However, most open-pollinated progeny were the result of xenogamy (outcrossing rate, tm=0.68). The study shows that floral contrivances and pollination system have a strong influence on mating pattern. The differential foraging behaviour of the pollinators causes deposition of a mixture of self- and cross-pollen to produce a mixed brood. Inbreeding depression and geitonogamy appear to play a significant role in sustaining mixed mating in this species.

Demography and mating system shape the genome-wide impact of purifying selection in Arabis alpina

Plant mating systems have profound effects on levels and structuring of genetic variation and can affect the impact of natural selection. Although theory predicts that intermediate outcrossing rates may allow plants to prevent accumulation of deleterious alleles, few studies have empirically tested this prediction using genomic data. Here, we study the effect of mating system on purifying selection by conducting population-genomic analyses on whole-genome resequencing data from 38 European individuals of the arctic-alpine crucifer Arabis alpina We find that outcrossing and mixed-mating populations maintain genetic diversity at similar levels, whereas highly self-fertilizing Scandinavian A. alpina show a strong reduction in genetic diversity, most likely as a result of a postglacial colonization bottleneck. We further find evidence for accumulation of genetic load in highly self-fertilizing populations, whereas the genome-wide impact of purifying selection does not differ greatly between mixed-mating and outcrossing populations. Our results demonstrate that intermediate levels of outcrossing may allow efficient selection against harmful alleles, whereas demographic effects can be important for relaxed purifying selection in highly selfing populations. Thus, mating system and demography shape the impact of purifying selection on genomic variation in A. alpina These results are important for an improved understanding of the evolutionary consequences of mating system variation and the maintenance of mixed-mating strategies.

Patterns of chasmogamy and cleistogamy, a mixed-mating strategy in an endangered perennial.

Cleistogamy (CL) in angiosperms historically has been understudied; however, its co-occurrence with chasmogamy (CH) across many plant species suggests a fitness advantage to maintaining this mixed-mating strategy. Maintenance of mixed-mating has been attributed to reproductive assurance, resource allocation or genetic trade-offs. Our goals were to explore patterns of CH and CL, quantify reproductive contributions measured by fruit production and determine how CL is maintained in the endangered perennial Polygala lewtonii. This species exhibits CH and both above-ground cleistogamy (CL-AG) and below-ground cleistogamy (CL-BG). In monthly censuses from 2008 to 2012, we documented flowering patterns by counting CH flowering stems, CL-AG fruits and CL-BG rhizomes per plant. Monitoring of buds on CH flowering stems in 2004 provided an estimate of CH fruits per plant. Plant excavations in 2005 of CL-BG rhizomes provided an estimate of CL-BG fruits per plant. Floral morphs were temporally separated with CH flowers observed from January to May and CL flowers from June to February. Overall, 17.5 % of plants flowered; most plants expressed CH first in spring months (63.4 %) and the rest initiated CL-AG in fall months. Reproductive output was dominated by CH (median 26 fruits) compared to combined CL (median 3.5 fruits). Annual reproductive effort of CL-AG was positively correlated with plant age while CH had no relation. Our research shows CH as the dominant form of reproductive effort with most individuals expressing CH and through greater reproductive contributions. CL appears limited by plant size or resources based on the positive relationship with plant age. CL dependency on resource availability is common in other species found in dry or low-quality habitats; however, CL contributions in this species are comparatively low. This raises more questions related to energy requirements of both floral morphs, how this affects the production of viable progeny and why CL persists.

Diversity, distribution, and significance of transposable elements in the genome of the only selfing hermaphroditic vertebrate Kryptolebias marmoratus

The Kryptolebias marmoratus is unique because it is the only self-fertilizing hermaphroditic vertebrate, known to date. It primarily reproduces by internal self-fertilization in a mixed ovary/testis gonad. Here, we report on a high-quality genome assembly for the K. marmoratus South Korea (SK) strain highlighting the diversity and distribution of transposable elements (TEs). We find that K. marmoratus genome maintains number and composition of TEs. This can be an important genomic attribute promoting genome recombination in this selfing fish, while, in addition to a mixed mating strategy, it may also represent a mechanism contributing to the evolutionary adaptation to ecological pressure of the species. Future work should help clarify this point further once genomic information is gathered for other taxa of the family Rivulidae that do not self-fertilize. We provide a valuable genome resource that highlights the potential impact of TEs on the genome evolution of a fish species with an uncommon life cycle.

Population genetics and evolution of the mangrove rivulus Kryptolebias marmoratus, the world's only self-fertilizing hermaphroditic vertebrate.

The mangrove rivulus, Kryptolebias marmoratus (Rivulidae, Cyprinodontiformes), is phylogenetically embedded within a large clade of oviparous (egg laying) and otherwise mostly gonochoristic (separate sex) killifish species in the circumtropical suborder Aplocheiloidei. It is unique in its reproductive mode: K. marmoratus is essentially the world's only vertebrate species known to engage routinely in self-fertilization as part of a mixed-mating strategy of selfing plus occasional outcrossing with gonochoristic males. This unique form of procreation has profound population-genetic and evolutionary-genetic consequences that are the subject of this review.

Mating system modulates degree of seed dormancy inHypericum elodesL. (Hypericaceae)

AbstractKnowledge of processes responsible for seed dormancy can improve our understanding of the evolutionary dynamics of reproductive systems. We examined the influence of the breeding system on primary seed dormancy inHypericum elodes, an Atlantic–European softwater pools specialist plant that exhibits a mixed mating strategy (the ability to both self- and cross-pollinate) to set seeds. Seeds were obtained through hand pollination treatments performed in a natural population during three consecutive years. Primary dormancy of seeds recovered from each pollination treatment was measured by analysing the seed germination response at dispersal and after various periods of cold stratification. While all collected seeds exhibited physiological dormancy, the degree of primary dormancy was associated with the pollination treatments. Weak and rapid loss of primary dormancy characterized seeds recovered from self-pollinated flowers, while stronger dormancy was found in seeds obtained from cross-pollination. The association between pollination treatments and primary dormancy indicated that the mating system should be considered as a source of variation for dormancy degree, proportional to self- and cross-pollinations (selfing rate) within populations of this species. These results suggest that by shedding seeds with various degrees of dormancy, plants may distribute their offspring across time by means of polymorphism in germination response. We conclude that seed germination alone is not an appropriate fitness measure for inbreeding depression estimates, unless dormancy is removed.