Fruit set increases with maleness in the andromonoecious Acacia caven

In long-lived plants, sexual expression can be highly variable in time and space. The extent to which sexual dimorphism of flowers is expressed in the same or in separate individuals, may have implications for the plants’ dependence on pollen vectors, the compatibility of the pollen received, and the potential for setting fruit. Here, we account for variability in sexual expression in the small tree Acacia caven, which produces hermaphrodite and male flowers on the same or in separate individuals. We focus on whether individual sexual expression influences fruit set, which latter has been reported to be extremely low in this species. Individual sexual expression, estimated as the hermaphrodite: male flowers ratio was variable both within and among plants across two reproductive seasons. Though male flower production could be very high (up to 99% of the flowers), all trees produced both flower types within the same individual, thus confirming andromonoecy in this species. More interestingly, hermaphrodite: male flowers ratio had a negative effect on fruit set. This pattern was consistent across two reproductive seasons and suggests that maleness should favor female function. Similar trends found in other plant species with the same sexual system support the hypothesis that male flowers of andromonoecious species may contribute to reduce self-pollen interference compared with hermaphrodite flowers. By favoring the deposition of compatible pollen grains and increasing fruit set, maleness would then help to maintain andromonoecy in this and other andromonoecious species.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

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.     

Floret specialization, seed production and gender in Artemisia vulgaris L. (Asteraceae, Anthemideae)

Flowering and fruiting behaviour of female and hermaphrodite florets is described and assessed in samples from three populations from Denmark, England and Sweden. Between 25 and 50% of the florets in capitula are female, and flowering gender varies little among plants in each population. Fruiting gender of individuals, G (femaleness), varies from 0 to 0.85, because of variation in fruit set and fruit abortion. Variation in fruiting gender was correlated with plant size parameters in two populations, but not in the third. The data suggest that post-anthesis regulation of maternal investment may be operating. Florets of A. vulgaris are either totally specialized for pollen receipt (female florets) or largely specialized for pollen donation (hermaphrodite florets), and show adaptations for avoiding interference with each other in these functions. Movement of capitula from a pendent position at flowering to an erect position at fruiting optimizes positions for dissemination of pollen and of seeds respectively.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation

 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002     

Reproductive characters in a gynodioecious species, Silene italica (Caryophyllaceae), with attention to the gynomonoecious phenotype

Gynodioecious populations (i.e. populations with female and hermaphrodite individuals) often contain a third phenotype with an intermediate sex expression. In Silene italica , this phenotype is characterized by a mixture of pistillate and perfect flowers and is thus gynomonoecious. To characterize sexual functions of these gynomonoecious individuals and their potential influence in the maintenance of gynodioecy, reproductive characters of the three sexual phenotypes were compared over 2 years in several families of S. italica produced by crossing. We found that gynomonoecious individuals were intermediate for flower production and female fertility characters, although they did not always significantly differ from female individuals. Perfect flowers of gynomonoecious and hermaphrodite plants were similar in size and pollen production. Gynomonoecious individuals were thus intermediate in female and in male functions. Family effects were found for most of the characters. The female advantage (i.e. the fertility of females compared to the female fertility of pollen producing plants) was not dramatically different when gynomonoecious plants were taken into account.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 583–591.     

Sex expression and fruit set in Silene littorea (Caryophyllaceae): variation among populations

The flowers of Silene littorea may be female or hermaphrodite, and an individual may show any of three sexual phenotypes: female-flowers-only, hermaphroditeflowers-only, or mixed-type. We estimated the relative frequencies and fruit sets of each flower type and each sexual phenotype in eleven S. littorea populations in northwest Spain. For all plants and sites combined, 18% of flowers were female and 82% hermaphrodite. Individuals either had only female flowers (6%), only hermaphrodite flowers (43%), or had both flower types (51%). The relative frequencies of female and hermaphrodite flowers, and of sexual phenotypes, vaned among populations. There was no significant effect of flower sex on its probability of setting fruit. Differences among individual plants within populations had the most prominent role in explaining variation among flowers in probability of setting fruit, while population and its interaction with flower sex had a negligible and nonsignificant influence. Mean fruit set per plant did not differ significantly between sexual phenotypes. Neither population nor its interaction with sexual phenotype accounted for significant amounts of between-plant variance in fruit set.     

冠果草的性表达状态及其进化含义

通常认为雄花两性花同株是植物性表达两性花进化到雌雄异株的过渡类型之一。慈姑属仅冠果草（Ｓａｇｉｔｔａｒｉａ ｇｕｙａｎｅｎｓｉｓ Ｈ．Ｂ．Ｋ．ｓｕｂｓｐ．ｌａｐｐｕｌａ（Ｄ．Ｄｏｎ）Ｂｏｊｉｎ）的性表达为典型的雄花两性花同株,其他种为雌雄同株。对冠果草野外和实验居群的性配置进行了定量观察。东乡和武夷山居群雄花的比例分别为２．４８％０．９６％。实验表明,栽培条件影响冠果草的性表达：在营养较好的条件下     

Effects of population size, seed predation and plant size on male and female reproductive success in Vincetoxicum hirundinaria (Asclepiadaceae)

Here we present the results of a study on the effects of population size, seed predation, and plant size on male and female reproductive success in ten Vincetoxicum hirundinaria populations in southwestern Finland. We investigated male reproductive success by recording pollinia removal, and studying pollen quality and quantity. Pod and seed production, and the number of ovules per ovary were counted to estimate female reproductive success. Both pollen quality and pollinia removal were higher in small V. hirundinaria populations compared to large populations. In contrast, the quantity of pollen was higher in large populations than in small populations. Pod initiation was higher in large populations. However, large populations had higher abortion rates and proportionally fewer intact pods. Plant size did not affect male reproductive success whereas pod and seed production correlated positively with plant size, although only in large populations. Population size had no clear impact on seed predation intensities. Variation was, however, observed among populations. Larger plant individuals suffered from an increased risk of being attacked only in two populations. This correlation was not clearly attributed to plant population size. The results of this study suggest that reproductive success in V. hirundinaria may be highly variable and this variation is to some extent explained by population size. Further, population size affected male and female reproductive success differently.     

Sex Expression and the Evolutionary Advantages of Male Flowers in an Andromonoecious Species, Sagittaria guyanensis subsp. lappula (Alismataceae)

Sex expression in Sagittaria guyanensis H.B.K. subsp. lappula (D. Don) Bojin is typically andromonoecious while the other species in the same genus are basically monoecious. The evolutionary advantages of male flowers and hermaphrodite flowers in S. guyanensis subsp. lappula were assessed by measuring sex allocation and pollen movements in two wild populations of the species. Two cultivated populations served as controls. The percentage of male flowers was very low in the two wild populations in Dongxiang, Jiangxi Province and Wuyishan, Fujian Province, viz., 2.48% and 0.96% respectively. In the two cultivated populations, male flower percentage significantly increased when the soil was of higher nutrient content. This indicates that the allocation to male versus female reproduction might change in response to environmental factors. Pollen production per male flower was 4.1 times higher than that of a hermaphrodite flower. The floral shape and size of male and hermaphrodite flowers were similar. No difference was observed between these dimorphological flowers in pollen germination rate in vitro and in the speed of pollen tube growth in vivo. Anthesis was only 4-5 h. Male flowers usually opened 0.5 h earlier than hermaphrodite flowers. An unexpected finding was that no pollen from the male flowers was found on the stigmas of the hermaphrodite flowers, in spite of the occasional visits by insects to both types of flowers in both wild and control populations. A consistent pattern of fruit development was found to exist in open pollinated flowers as well as in flowers that had been bagged. The sex ratios did not have significant influence on fruit set. Approximately 25% of the pistils in a gynoecium failed to develop into fruits because no pollen was deposited on them, indicating that the fruit set of this andromonoecious plant is mainly affected by pollen limitation rather than resource limitation. Reproduction in S. guyanensis subsp. lappula in the habitats was dependent on self-pollination in hermaphrodite flowers. The male flowers in this species might be a potential source of additional pollination and may facilitate cross-pollination. The fact that the flowers of monoecious species in Sagittaria pollinated by a wide diversity of insect visitors may contribute enormously to the diversification of sex expression in this genus.     

Population size,pollinator visitation and fruit production in the deceptive orchid Calypso bulbosa

It has been proposed that in non-rewarding animal-pollinated plants the pollination intensity should decrease with increasing population size and should increase with increasing local abundance of reward-producing plants. To test these hypotheses, we examined how population size, local abundance of Salix caprea, and tree cover were related to pollen removal and fruit production in 16 populations of the deceptive, early-flowering and bumblebee-pollinated orchid Calypso bulbosa in northern Sweden in 3 consecutive years. To determine whether fruit production was limited by pollinator visitation, supplemental hand-pollinations were performed in three populations in 3 years. Finally, to examine whether increased fruit production was associated with a reduction in future flower production, vegetative growth or survival, supplemental hand-pollination was repeated for 5 years in one population. The levels of pollen export, pollen deposition, and fruit set of C. bulbosa varied considerably among years and among populations. The proportion of plants exporting pollen was negatively related to population size, and positively related to density of S. caprea and to tree cover in 1 of the 3 years. In the other 2 years, no significant relationship was detected between proportion of plants exporting pollen and the latter three variables. In no year was there a significant relationship between fruit set and population size, density of S. caprea and tree cover. There was substantial among-year variation in the extent to which fruit production was limited by insufficient pollen deposition and in the amount of weather-induced damage to flowers and developing fruits. Fruit set was consistently higher in hand-pollinated than in open-pollinated plants, but this difference was statistically significant in only one of 3 years. Supplemental hand-pollination in 5 consecutive years increased cumulative fruit production 1.8 times, but did not affect flower production, plant size, or survival. Tree cover was negatively correlated with the incidence of frost damage in 1 year. The results indicate that life-time seed production may be pollen limited in C. bulbosa, and that variation in population size and local abundance of the early-flowering, nectar-producing S. caprea can only partly explain the extensive variation in pollinator visitation among populations of this species.     

白背叶(大戟科)性系统和传粉生物学研究

为探讨野桐属(Mallotus)雌雄异株的进化和传粉机制,对白背叶(Mallotus apelta)野生居群的性系统和传粉生物学进行了研究。结果表明,所调查的白背叶居群均由雌性(F)、雄性(M)和少数两性(B)个体组成,平均性比为1：0.66：0.18(F：M：B);3种性别植株的基径大小差异不显著;雌株与两性植株的单花胚珠数、单果结籽数和自然结实率无显著差异;雄株与两性植株的花粉萌发率、花粉组织化学和花粉微形态特征也无显著差异,但雄株的单花花粉量是两性植株的1.26~1.63倍,且差异显著;雌株的异交结实率为96.67%,两性植株的异交结实率为76.00%,两者差异显著,说明居群内雌株的潜在种子生产力明显高于两性植株;野外观察到雄株和两性植株上的雄花具有访花者而雌株没有;雌株经套网处理后结实率超过30%而套袋处理不结实。这些表明白背叶具典型的亚雌雄异株性系统,雌株和雄株的适合度均高于两性植株;雌株以风媒传粉结实,两性植株可能兼有风媒和虫媒传粉特征。     

Effects of pollen reward removal on fecundity in a self-incompatible hermaphrodite plant

The pollen of hermaphrodite plants is often utilised by flower-visiting animals. While pollen production has obvious benefits for plant male fitness, its consequences for plant female fitness, especially in self-incompatible hermaphrodite species, are less certain. Pollen production could either enhance seed production though increased pollinator attraction, or reduce it if ovules are discounted by deposition of self pollen, as can occur in species with late-acting self-incompatibility. To test the effects of pollen reward provision on female fitness, we artificially emasculated flowers in two populations of the succulent Aloe maculata (Asphodelaceae), which has a late-acting self-incompatibility system, over the course of its flowering period. Flowers of this species are visited by sunbirds (for nectar) and native bees (for pollen and nectar). We measured floral visitation rates, floral rejection rates, pollen deposition on stigmas and fruit and seed set in both emasculated and non-emasculated plants. We found that flowers of emasculated plants suffered reduced visitation and increased rejection (arrival without visitation) by bees, but not by sunbirds; had fewer pollen grains deposited on stigmas and showed an overall decrease in fruit set and seed set. Rates of seed abortion were, however, greatly reduced in emasculated flowers. This study shows that pollen rewards can be important for seed set, even in self-incompatible plants, which have been assumed to rely on nectar rewards for pollinator attraction. Seed abortion was, however, increased by pollen production, a result that highlights the complexity of selection on pollen production in hermaphrodite flowers.     

Scale-specific determinants of a mixed beech and oak seedling–sapling bank under different environmental and biotic conditions

Based on the hypothesis that both plant size and local conspecific density influence allocation to female/male functions, we explored the relationship between plant height, local conspecific density, sexual expression, and fruit production in the andromonoecious shrub Caesalpinia gilliesii. We quantified the total number of perfect and staminate flowers, the pollen received and fruits produced per plant in two populations, and estimated phenotypic gender and fruit set. Local density failed to explain phenotypic gender, nevertheless, plant height and fruit set increased with local density in one population where, in addition, the slopes for the size-dependent sex allocation curve were steeper. As observed for other plant species, this suggests that between population differences in resource availability is the main underlying factor for the observed population differences in the size-dependent allocation pattern to flowers and fruits. On the other hand, the number of staminate and perfect flowers per plant increased with plant height and the fastest increase of staminate flowers resulted in a male-biased size-dependent sex allocation strategy in both populations. Since pollination intensity was not correlated with plant height in any population, the observed allocation strategy cannot be attributed to differences in pollen availability between different sized individuals, but to differences in plant size. Finally, because fruit set and total fruit number increased with plant height in one population, the obtained results provide further evidence that animal-pollinated, andromonoecious species may exhibit a male-biased size-dependent sex allocation strategy, which may favor female fecundity.     

湿地植物大花百合种群的性系统特征

开花植物性系统特征是植物在长期适应进化过程中逐渐形成的繁殖策略, 它在一定程度上影响了物种的种群结构及其在生境中的分布格局, 因此性系统特征一直是植物进化与生态学领域的研究热点。本研究于2012年和2020年在吉林省金川泥炭沼泽湿地, 通过样地调查取样的方法, 研究了湿地植物大花百合(Lilium concolor var. megalanthum)的性系统特征以及不同性别表型植株的比例、密度及空间分布格局, 并探究了大花百合植株个体大小和其性别表达的联系, 比较了不同性别表型的花粉实际可育性和来源不同的花粉授粉处理结实后的种子活力。结果显示, 个体水平上, 大花百合具有雄花植株、两性花植株和雄花两性花同株(即雄全同株) 3种性表型; 种群水平上, 具有雄花的个体充当父本, 通过花粉向下一代传递基因, 而具有两性花的个体主要充当母本, 通过胚珠实现基因的传递。2020年大花百合雄花植株占种群内开花植株的39.14%, 相比2012年雄花植株比例增加了22.80%; 大花百合种群分布密度为0.06株/m², 较2012年下降了0.09株/m²。从2012年到2020年, 总体表现为种群密度降低, 雄花植株相对比例增加的趋势。两性花植株和雄花植株在小尺度范围主要呈聚集分布, 随着尺度增大, 趋于随机分布。雄花植株显著小于两性花植株, 但鳞茎资源分配显著高于两性花植株。大花百合植株的性表达遵循大小依赖的性分配假说: 个体较小的植株表达为雄花植株, 而个体较大的植株则为两性花植株或雄全同株。在不同授粉处理下的结籽率和种子萌发结果表明, 大花百合自花授粉结籽率和坐果率显著低于异花授粉, 雄花植株和两性花植株花粉可育性以及对两性花植株授粉后所获得种子活力无显著差异。大花百合雄花植株的出现可能是对有限环境资源条件适应的结果, 反映了湿地条件下该植物在生长和繁殖资源之间的权衡策略, 在长期的湿地演变过程中, 这种性系统特征的出现具有一定的适应性意义。     

Size-dependent sex allocation in a monocarpic perennial herb, <Emphasis Type="Italic">Cardiocrinum cordatum</Emphasis> (Liliaceae)

The size-dependent sex allocation model predicts that the relative resource allocation to female function often increases with plant size in animal-pollinated plants. If size effects on reproductive success vary depending on the environmental conditions, however, the size dependency may differ among populations. We tried to detect site-specific variation in size-dependent sex allocation of a monocarpic hermaphrodite with reference to light availability. Multiple flowers and fruits were sampled from the individuals of Cardiocrinum cordatum, a monocarpic understory herb, and pollen, ovule and seed production were measured with reference to the plant size in two populations. Furthermore, frequency and foraging behavior of pollinator visitation was observed. Ovule production per flower increased with plant size in both populations, while pollen production per flower increased with size only in the population under sparse canopy. Therefore, proportional allocation to male function decreased with plant size in the population under closed canopy, but did not change in the population under sparse canopy. Pollinators usually visited only one flower per plant, indicating the negligible geitonogamous pollination in this species. Although seed production under closed canopy was lower than that under sparse canopy, seed-set rate per flower and seed mass per fruit were independent of plant size in either of the populations. Size-dependent sex allocation in this species was site-specific, suggesting that not only resource storage before reproduction (i.e., plant size) but also resource availability of environment throughout the reproductive process (i.e., light availability) affect reproductive performance in this species.     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.     

Pollen Production in the Androdioecious Datisca glomerata (Datiscaceae): Implications for Breeding System Equilibrium

Abstract Androdioecy (the presence of males and hermaphrodites in a breeding population) is a rare reproductive system in plants, with Datisca glomerata (Datiscaceae) representing the only well-documented example. Recent reports of high outcrossing rates, inbreeding depression, and high male pollen production satisfy theoretical predictions for the continued maintenance of androdioecy in populations of this species. However, in prior studies pollen production was measured indirectly in terms of numbers of anthers per flower—based on the assumption that male and hermaphroditic plant have equal numbers of flowers and that anthers from the two sexual morphs produce equivalent amounts of pollen. Herein, we demonstrate that male and hermaphrodite plants do not differ significantly in terms of flower number, but that pollen production in anthers from hermaphroditic plants is 12.6% higher than in anthers from male plants, thus refining the estimate of relative pollen fecundity of male versus hermaphrodite plants. The differential lowers the frequency of males predicted by theory, but is still consistent with the maintenance of androdioecy in this species.     

Positive effects of flower abundance and synchronous flowering on pollination success,and pollinia dispersal in rewardless Changnienia amoena (Orchidaceae)

Pollination success and pollen dispersal in natural populations depend on the spatial‐temporal variation of flower abundance. For plants that lack rewards for pollinators, pollination success is predicted to be negatively related to flower density and flowering synchrony. We investigated the relationships between pollination success and flower abundance and flowering synchrony, and estimated pollinia dispersal distance in a rewardless species, Changnienia amoena (Orchidaceae). The results obtained in the present study revealed that male pollination success was negatively influenced by population size but was positively affected by population density, whereas female pollination success was independent of both population size and density. Phenotypic analysis suggested that highly synchronous flowering was advantageous through total pollination success, which is in contrast to previous studies. These results indicate that pollination facilitation rather than competition for pollinator visits occurs in this rewardless plant. The median distance of pollinia dispersal was 11.5 m (mean distance = 17.5 m), which is comparable to that of other rewardless plants but longer than for rewarding plants. However, pollen transfer occured mainly within populations; pollen import was a rare event. Restricted gene flow by pollinia and seeds probably explains the previous population genetic reporting a high degree of genetic differentiation between populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 477–488.