SEX RATIO,SEED PRODUCTION,BIOMASS ALLOCATION,AND THE COST OF MALE FUNCTION IN CUCURBITA FOETIDISSIMA HBK (CUCURBITACEAE)

In the gynodioecious plant Cucurbita foetidissima (Cucurbitaceae), females were common in all eight populations examined and made up 32% of adult plants. Females produced 1.5 (SE = 0.2) times as many seeds as did hermaphrodites. The observed difference in seed production alone is not great enough to explain the maintenance of females, especially at their current frequency. Females and hermaphrodites did not differ in number of nodes per stem, stems per plant, internode length, or size of leaves. Females produced more female biomass (fresh or dry weight) than hermaphrodites, but total investment in sexual biomass did not differ. Thus, the biomass of male flowers produced by hermaphrodites was about equal to the extra female biomass produced by females. The results support the existence of a trade-off between male and female reproduction.     

The functional ecology of gynodioecy in Eritrichum aretioides (Boraginaceae), the alpine forget-me-not

Eritrichum aretioides is a gynodioecious species with female and hermaphrodite individuals. In populations on Pennsylvania Mountain in central Colorado (USA), the frequency of females ranges from 22 to 41%. Flower number and the number of seeds produced per flower were similar in hermaphrodites and females. However, hermaphrodites produced larger flowers, while females produced larger seeds (P < 0.05 for both). In the field, seed germination was higher for seeds from females than for seeds from hermaphrodites (20 vs. 9% germination; P < 0.05). Unvisited flowers and open-pollinated flowers of hermaphrodites had similar pollen receipt (approx 20 pollen grains per stigma), but seed set following autogamous pollination was significantly lower than seed set following natural pollination. This finding indicates that hermaphrodites have a barrier to selfing and implies that the larger seed size and greater establishment advantage of offspring from females is unlikely to have resulted from female outcrossing advantage. Rather, differences in the quality of seed progeny between morphs probably reflect a trade-off in sexual allocation or pleiotropic effects of the sex-determining genes.     

The first record of gynodioecy in a species of Gnidia (Thymelaeaceae) from South Africa

Sexual polymorphism was studied in the shrub Gnidia wikstroemiana (Thunb.) Meisn. from the semiarid Nama Karoo Biome, South Africa. The populations comprised plants bearing either female flowers, or hermaphrodite flowers with variable female function. In two populations, female plants accounted for 36–37% of the flowering plants. Female flowers were smaller and their stamens were reduced to staminodes, but their styles were significantly longer than those of hermaphrodite flowers. Energy investment in flowers and fruits for females and hermaphrodites was measured using bomb calorimetry. Females produce a greater number of less costly flowers than hermaphrodites, and invest less energy per unit in production of flowers and inflorescences. In contrast, females invest more energy per unit in production of fruits and infructescences than hermaphrodites. Females overall invest 7.3% more energy in reproduction than hermaphrodites. Female flowers were obligate out-crossers (xenogamous), with 35% of nonmanipulated, open-pollinated flowers setting fruit, comparable with fruit set among selfed hermaphrodite flowers. The breeding strategy of G. wikstroemiana most closely resembles gynodioecy. This is the first report of sexual dimorphism in Gnidia L. and sub-Saharan Thymelaeaceae.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 219–233.     

RESOURCE COMPENSATION AND THE EVOLUTION OF GYNODIOECY IN PHACELIA LINEARIS (HYDROPHYLLACEAE)

Gynodioecy is a dimorphic breeding system in which hermaphrodite and female individuals coexist in populations. Theoretical models have shown that if nuclear genes control sex expression, then gynodioecy can evolve only when females have large advantages in one or more fitness components. These female advantages must be large enough that females' expected lifetime production of viable seeds is more than twice that of hermaphrodites. Previous studies have found that cytoplasmic inheritance and/or a large offspring-vigor advantage of females (caused by hermaphrodite self-pollination and inbreeding depression of selfed seeds) account for this breeding system's evolution. This paper reports studies of gynodioecy in Phacelia linearis, an insect-pollinated annual plant in which gender inheritance appears to be nuclear. Twenty-six P. linearis populations surveyed in northern Utah, USA, contain a majority of perfect-flowered hermaphrodites, but most (22) also contain male-sterile individuals (females), at frequencies of up to 0.16. The hermaphrodite selfing rate is low (0.00–0.20 in four populations). Maternal gender does not consistently affect components of offspring vigor, such as seed size, germination rate, seedling survivorship, and vegetative size. Plants of the two genders do not differ in number of seeds per fruit or mean seed mass. Females produce significantly more fruits and seeds than hermaphrodites in natural populations. The ratio of the mean lifetime seed production of females to the mean lifetime seed production of hermaphrodites ranged from 1.31 to 2.52 in six natural populations. Females have greater shoot biomass than hermaphrodites and produce more seeds at any given shoot biomass than hermaphrodites, suggesting that their seed-production advantage arises from gender-specific patterns of resource allocation to growth and reproduction. The gender difference in plant size varies across environments and across genetic backgrounds. In this species nuclear gynodioecy appears to be evolutionarily stable mainly because of resource compensation by females, without a large outcrossing advantage of females.     

Maintenance of gynodioecy in Wurmbea biglandulosa (Colchicaceae): gender differences in seed production and progeny success

 In gynodioecious species, females contribute genes to future generations only through ovules, and to persist in populations they must have a compensatory advantage compared with hermaphrodites that reproduce via ovules and pollen. This compensation can result from greater fecundity and/or superior success of progeny from females. We examined differences in seed production and progeny success between females and hermaphrodites in the geophyte Wurmbea biglandulosa to explain the maintenance of females. Females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites but this did not necessarily result in greater fecundity, in part because seed production of females was pollen-limited. Over four years in one population, open-pollinated females produced 1.32 more seeds than open-pollinated hermaphrodites (range 1.09–1.63). In two other populations examined for one year only females produced 1.07 and 0.79 as many seeds as hermaphrodites. Seed production of open-pollinated females and hermaphrodites was only 55% and 73% that of cross-pollinated plants, respectively, indicating that both genders were pollen-limited but females more so than hermaphrodites. Open-pollinated seeds from females were 1.18–1.27 times more likely to germinate than seeds from hermaphrodites. No gender differences existed in seedling growth or survival. Hermaphrodites were self-compatible, but selfed seed set was only 80% that of crossed seed set. Crossed seed set of females and hermaphrodites did not differ. Assuming nuclear control of male sterility, relative female fitness is insufficient to maintain females at their current frequencies of 17%, and substantial female fitness advantages at later life-cycle stages are required. Received May 4, 2001 Accepted February 25, 2002     

THE RELATIVE IMPORTANCE OF INBREEDING AND MATERNAL SEX IN DETERMINING PROGENY FITNESS IN SIDALCEA OREGANA SSP. SPICATA,A GYNODIOECIOUS PLANT

Gynodioecious plant populations contain both hermaphrodite and female individuals. For females to be maintained they must compensate for their loss of reproductive fitness through pollen. Females may achieve compensation by producing more and/or higher quality seeds than hermaphrodites. In this study, I investigated the independent and interactive effects of maternal sexual identity and inbreeding level on fitness of the progeny of hermaphrodites and females of Sidalcea oregana ssp.spicata. Seeds produced by selling hermaphrodites and by outcrossing or sib-crossing hermaphrodites and females, were planted in the field and greenhouse. Maternal-sex effects were substantial at the juvenile stages of the life cycle; seeds of females germinated in higher proportions and produced seedlings that grew significantly faster. Inbreeding effects were manifested primarily at the adult stage of the life cycle. Outcrossed plants were significantly larger and produced more flowers per plant than sib-crossed and selfed plants growing in the greenhouse. Progeny of hermaphrodites and females appeared to respond similarly to sib-matings. The maternal-sex effects observed in Sidalcea may have been related to cytoplasmically inherited factors and could be a driving force in the maintenance of females. Inbreeding depression could play a role in determining the fitness of both sex morphs, if females experience biparental inbreeding in the field. Frequent inbreeding of hermaphrodites may not be necessary to explain the maintenance of gynodioecy in this species.     

Female frequency and relative fitness of females and hermaphrodites in gynodioecious Geranium sylvaticum (Geraniaceae)

We determined female frequency of 23 populations of the gynodioecious Geranium sylvaticum (Geraniaceae) in Finland. We compared our results to previous results on this species from the 1960s in order to reveal putative changes in female frequencies. Because females may be maintained in gynodioecious populations if their seed production or offspring quality is higher than that of hermaphrodites, we explored reproductive success of females and hermaphrodites in detail in 11 populations for two consecutive years. Female frequencies varied from 0.4 to 27.2%; this variation is similar to that observed in the 1960s. Contrary to previous results that indicated lower seed production in females, females produced 1.2 and 1.7 times more seeds per flower than hermaphrodites in 2000 and 2001, respectively. Females also had higher fruit set than hermaphrodites. Thus, higher seed production of females partly explains the maintenance of gynodioecy in this species. Furthermore, female frequency correlated negatively with relative seed fitness of hermaphrodites suggesting that relative seed fitness is related to population sex ratio. Female frequency and the distance of the population from the most southern population also tended to correlate positively, suggesting that harsher environmental conditions in the north may benefit female plants. Given the observed yearly variation, our results also highlight the importance of temporal variation for the relative seed fitness of females and hermaphrodites.     

Population Size,Pollination and Breeding System of Silene stockenii Chater (Caryophyllaceae), an Annual Gynodioecious Species of Southern Spain

The distribution area, phenology, sex polymorphism, floral characteristics and breeding system of Silene stockenii (Caryophyllaceae), a narrowly endemic annual species of southern Spain, were studied. Only five populations were found in a total area of 2 ha. Silene stockenii is a gynodioecious species with fully female, fully hermaphrodite and intermediate plants bearing hermaphrodite, male-sterile, and partially male-sterile flowers. Male-sterile flowers are typically smaller than hermaphrodites. Nectar production was significantly higher in hermaphrodite plants and during the female phase of hermaphrodite flowers. The red flowers appear during the spring (March-May) and are pollinated by long-tongued Bombyliidae. Hand pollinations revealed that the species is self-compatible; however, natural self-pollination is rare due to marked protandry. Hand pollination significantly increased the number of seeds per fruit and seed set, indicating limited pollination in the field. In controlled pollinations female plants of S. stockenii produced higher seed set than hermaphrodite plants, but in freely pollinated plants fruit set and seed production was similar in both morphs, indicating that pollinators do not discriminate in favour of hermaphrodite plants.     

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.     

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.     

Sex‐specific reproductive components and pollination ecology in the subdioecious shrub Fuchsia microphylla

In subdioecious populations, functional female, male and hermaphrodite individuals coexist. Subdioecy may be a transitional state towards dioecy or a breakdown of dioecy, although lability in sex expression may maintain subdioecy as a stable condition. To better understand the ecological aspects involved in sex ratio dynamics and breeding system evolution, we studied the pollination and female fitness components of female and hermaphrodite individuals of the subdioecious shrub Fuchsia microphylla. In two natural populations at the Trans‐Mexican Volcanic Belt we estimated female frequency and several reproductive components of female and hermaphrodite plants under natural pollination and experimental pollination treatments. Average female frequency was 42%, and on average, 42.5% of hermaphrodites produced fruits. Female plants showed a 17‐fold female fertility advantage over hermaphrodites through increased fruit production, as the number of seeds and germination rates did not differ between morphs. Hermaphrodite flowers were larger, with similar nectar production and concentration to female flowers, and pollinators did not show consistent morph preferences. Some hermaphrodites produced fruits autonomously, and female flowers excluded from pollinators produced fruits putatively by apomixis. Fruit production in hermaphrodites, but not in females, was related to height, suggesting increased investment of hermaphrodites in the female function at higher resource status. For sex ratios to be at equilibrium, the female fertility advantage should be reduced about eightfold. However, it may be that hermaphrodites are maintained by producing fruits at no cost to the male function at higher resource status, as the gender plasticity hypothesis proposes.     

Pre-dispersal seed predation in gynodioecious Geranium sylvaticum is not affected by plant gender or flowering phenology

Sex-specific interactions with antagonists may explain female maintenance in gynodioecious populations if seeds produced by hermaphroditic plants are preferred over seeds produced by female plants. Among antagonistic interactions, pre-dispersal seed predators have received relatively little attention even though they may exert sex-specific selective pressures on the evolution of floral and flowering traits. In this work, I investigate temporal variation in seed predation in gynodioecious Geranium sylvaticum, where in addition to female and hermaphrodite individuals, plants with an intermediate sexual expression are also present in most populations. Specifically, I examined whether seed predation is linked to flowering phenology, plant gender, and sexual dimorphism in floral and seed traits over the flowering season using an experimental field population. Within the population, I selected female, intermediate, and hermaphrodite plants with different timing of flowering onset (early, mid, or late), and collected seeds across the fruiting period. Seeds were weighed and examined for seed predator damage. The results show that the three genders experienced similar levels of seed predation attack regardless of their flowering phenology, and that overall seed predation was not related to changes in seed production or seed mass. These results suggest that sexual dimorphism in seed predation cannot be responsible for female maintenance in this species.     

Female compensation through the quantity and quality of progeny in a gynodioecious plant, Geranium maculatum (Geraniaceae)

One of the major evolutionary trends in flowering plants is the evolution of unisexual flowers (male or female) from perfect flowers. This transition has occurred repeatedly in many taxa and has generated a wonderful array of variation in sexual expression among species. Theoretical studies have proposed a number of mechanisms to explain how this level of variation could be maintained in natural systems. One possible mechanism is the female compensation hypothesis, which predicts that female mutants require an increase in their seed fitness in order to invade a hermaphroditic system. Using Geranium maculatum, I tested this hypothesis and showed that female mothers produced more and larger seeds than hermaphroditic mothers even though they were indistinguishable in their vegetative traits and the flower production. Seeds from females were also more likely to germinate and produced seedlings with larger above- and belowground biomass. These seedlings were more likely to flower than those from hermaphrodites in at least one of the two populations studied. Combined, these results indicated that females in G. maculatum did compensate for their loss of male function by producing more and better seeds than hermaphrodites. This provides a mechanism for the maintenance of female plants in this species.     

The role of pollination level on the reproduction of females and hermaphrodites in the gynodioecious plant Gypsophila repens (Caryophyllaceae)

In gynodioecious plant species, females are expected to have more resources available for maturing seeds because pistillate flowers are smaller, do not produce pollen, and are thus less costly that perfect flowers. The potential female advantage arising from more abundant resources is, however, likely to vary depending on whether seed production is limited by resource or pollen availability. Here we experimentally investigated the influence of pollen and resource limitation on female advantage in a gynodioecious species using two levels of pollination. Total seed production of females was always greater than that of hermaphrodites: females produced more flowers and more fruits that contained similar numbers of seeds of similar mass. Under low pollination, female and hermaphrodite plants allocated resources to increased flower production rather than to increased seed size or quality. We did not detect any influence of pollen or resource limitation on female advantage, which remained similar under low (= abundant resources) and full pollination. Outcrossed fruits performed better than selfed fruits when the same plant received both selfed and outcrossed pollen on different flowers. These differences were not greater under high pollination, possibly because resources available for each fruit did not differ between our pollen intensity treatments.     

Male‐biased hermaphrodites in a gynodioecious shrub,Daphne jezoensis

Gynodioecy, a state where female and hermaphrodite plants coexist in populations, has been widely proposed an intermediate stage in the evolutionary pathway from hermaphroditism to dioecy. In the gynodioecy–dioecy pathway, hermaphrodites may gain most of their fitness through male function once females invade populations. To test this prediction, comprehensive studies on sex ratio variation across populations and reproductive characteristics of hermaphrodite and female phenotypes are necessary. This study examined the variation in sex ratio, sex expression, flower and fruit production and sexual dimorphism of morphological traits in a gynodioecious shrub, Daphne jezoensis, over multiple populations and years. Population sex ratio (hermaphrodite:female) was close to 1:1 or slightly hermaphrodite‐biased. Sex type of individual plants was largely fixed, but 15% of plants changed their sex during a 6‐year census. Hermaphrodite plants produced larger flowers and invested 2.5 times more resources in flower production than female plants, but they exhibited remarkably low fruit set (proportion of flowers setting fruits). Female plants produced six times more fruits than hermaphrodite plants. Low fruiting ability of hermaphrodite plants was retained even when hand‐pollination was performed. Fruit production of female plants was restricted by pollen limitation under natural conditions, irrespective of high potential fecundity, and this minimised the difference in resources allocated to reproduction between the sexes. Negative effects of previous flower and fruit production on current reproduction were not apparent in both sexes. This study suggests that gynodioecy in this species is functionally close to a dioecious mating system: smaller flower production with larger fruiting ability in female plants, and larger flower production with little fruiting ability in hermaphrodite plants.     

COMPARATIVE REPRODUCTIVE BIOLOGY OF HERMAPHRODITIC AND MALE-STERILE IRIS DOUGLASIANA HERB. (IRIDACEAE)

A comparative study of the reproductive biology of male-sterile and hermaphroditic plants in a gynodioecious population of Iris douglasiana Herb. (Iridaceae) was conducted at the University of California's Marine Laboratory at Bodega Bay, California, between 1976–1979. Each year of the study, there were 11.1% male-sterile plants in the population, some of which began blooming at the same time as the earliest blooming hermaphrodites. Male-sterile flowers made up between 7–21% of the flowers produced during the male-sterile flowering period. Male-sterile flowers had smaller sepals and petals than hermaphrodites, there were fewer of them per square meter, and they had fewer pollinated stigmas than did hermaphroditic flowers. In a test to determine pollinator preference, intact hermaphroditic flowers tended to have more pollinated stigmas than did hermaphrodites with their stamens removed or those flowers with shortened sepals made to resemble the smaller male-sterile flowers. Floral phenology and nectar-flow patterns were similar in both types of flowers as were the kinds of amino acids and sugar rewards in the nectar. Male-sterile flowers, however, produced much less nectar per flower. There were no significant differences in the number of ovules per flower or the number of seeds produced per capsule between the two flower types, but the loss of seeds through larval predation was much greater in capsules from hermaphroditic flowers. Early flowering and setting of seed by plants with male-sterile flowers could give them a reproductive advantage over plants with hermaphroditic flowers which experience higher levels of larval predation later in the growing season.     

Sex-specific effect of Microbotryum violaceum (Uredinales) spores on healthy plants of the gynodioecious Gypsophila repens (Caryophyllaceae)

Females of gynodioecious species need to have reproductive advantages to compensate for their lack of male function and assure their maintenance in natural populations. Females may be more fecund than hermaphrodites because they reallocate resources from male to female function, avoid selfing and inbreeding depression, or produce higher quality offspring by screening arriving pollen better. A novel type of female advantage is proposed that may be important for several species of gynodioecious Caryophyllaceae. The anther smut fungus Microbotryum violaceum generally sterilizes its infected hosts but even without infection, spore deposition can negatively affect the reproduction of healthy individuals. In Gypsophila repens, flowers of hermaphrodite plants experimentally pollinated with both fungal spores and pollen produced significantly fewer fruits and seeds than those that received pollen alone, whereas female reproduction was unaffected by spore deposition. This unexplored reproductive advantage of females is probably due to the larger stigmatic surface in pistillate flowers, that allows pollen germination despite the presence of spores. Because longer stigmas may also lead to increased spore deposition, these results raise questions regarding the infection probability for each sex, possible sex-specific defense mechanisms, and the general role of pathogens in the maintenance of females in natural gynodioecious populations.     

Reproductive biology and relative male and female fitness in a trioecious cactus,Pachycereus pringlei (Cactaceae)

We describe the breeding system of an autotetraploid trioecious cactus, Pachycereus pringlei, provide estimates of the fitnesses of males and females relative to that of hermaphrodites, and discuss the role played by pollinators in the maintenance of three sexual morphs. Relatively high frequencies of females (45%) and males (26%) exist in coastal desert populations around Bahia Kino, Sonora, Mexico. They differ from hermaphrodites in flower size (females only), initiation of the flowering season, number of flowers produced per night and per season, sucrose content of nectar, and, in females, number of fruits produced per season under open pollination and in response to hand-pollination. Major similarities between the sex classes include overall plant size, nectar volume per flower, percent fruit set in open-pollinated flowers of females and hermaphrodites, seed mass and number of seeds per fruit, and pollen mass per flower in males and hermaphrodites. Hermaphrodites are self-compatible, and the selfing rate is high (65%). Levels of inbreeding depression in selfed fruits and seeds appear to be low. Fruit set is strongly pollinator-dependent in females but much less so in hermaphrodites. Relative fitness of males and females, as measured by annual production of pollen or seeds, is at least 1.5 times higher than that of the corresponding sex function in hermaphrodites. Given the high selfing rate and apparent lack of inbreeding depression, these fitness differences are insufficient to explain the occurrence of trioecy in this species.     

Males are cheaper,or the extreme consequence of size/age‐dependent sex allocation: sexist gender diphasy in Fritillaria montana (Liliaceae)

The bulbous geophyte Fritillaria montana is partially self‐compatible and capable of switching gender. Small flowering plants produce only single male flowers, but larger plants produce hermaphrodite or, rarely, male and hermaphrodite flowers. Eight populations in peninsular Italy were sampled to determine the frequency and size distributions of male and hermaphrodite plants, and to determine the relationship of plant size to male and hermaphrodite flower production. Male plants were significantly smaller than hermaphrodites and made up 14.5–47.8% (100% in one small population) of flowering plants within populations. There were no significant differences in male fitness among female‐sterile and hermaphrodite flowers, as they both possessed full and comparable fertilizing power. Therefore, the gender variation observed in F. montana is likely to depend on resource‐dependent sex allocation. From an evolutionary perspective, we highlight the occurrence of similar mechanisms of gender variation in other representatives of the order Liliales. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 323–333.     

Ecological context of breeding system variation: sex, size and pollination in a (predominantly) gynodioecious shrub

BACKGROUND AND AIMS: Species that exhibit among-population variation in breeding system are particularly suitable to study the importance of the ecological context for the stability and evolution of gender polymorphism. Geographical variation in breeding system and sex ratio of Daphne laureola (Thymelaeaceae) was examined and their association with environmental conditions, plant and floral display sizes, and pollination environment in a broad geographic scale was analysed. METHODS: The proportion of female and hermaphrodite individuals in 38 populations within the Iberian Peninsula was scored. Average local temperature and precipitation from these sites were obtained from interpolation models based on 30 years of data. Pollination success was estimated as stigmatic pollen loads, pollen tubes per ovule and the proportion of unfertilized flowers per individual in a sub-set of hermaphroditic and gynodioecious populations. KEY RESULTS: Daphne laureola is predominantly gynodioecious, but hermaphroditic populations were found in northeastern and southwestern regions, characterized by higher temperatures and lower annual precipitation. In the gynodioecious populations, female plants were larger and bore more flowers than hermaphrodites. However, due to their lower pollination success, females did not consistently produce more seeds than hermaphrodites, which tends to negate a seed production advantage in D. laureola females. In the northeastern hermaphroditic populations, plants were smaller and produced 9-13 times fewer flowers than in the other Iberian regions, and thus presumably had a lower level of geitonogamous self-fertilization. However, in a few southern populations hermaphroditism was not associated with small plant size and low flower production. CONCLUSIONS: The findings highlight that different mechanisms, including abiotic conditions and pollinator service, may account for breeding system variation within a species' distribution range and also suggest that geitonogamy may affect plant breeding system evolution.