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.     

Gynodioecy in Cirsium chikushiense Koidz. (Compositae)

A study of the morphology and function of flowers in Cirsiumchikushiense revealed that the species was distinctly gynodioecious.Self-incompatible hermaphrodite florets produced both seedsand pollen grains, while female ones produced seeds but no pollengrains at all. The degenerated stamens of females were not onlysmaller but also sometimes occurred at a lower position insidethe floral tube than in hermaphrodites. The stigmata of femalesoften developed more fully than those of hermaphrodites. Thefrequencies of female plants in natural populations varied from15·5 to 50%. Almost all the pollinators stayed on bothfemale and hermaphrodite heads only to collect nectar. The femaleplants of this species may be more specialized in their genderby saving the cost of not only pollen grains but also stamens,and may be maintained by sufficient pollinators in natural populations.This gynodioecy may provide an example of nuclear-cytoplasmicmale sterility.Copyright 1994, 1999 Academic Press Sex expression, gynodioecy, Cirsium chikushiense, Compositae, male sterility, degenerated stamen, female frequency     

Reproductive ecology of Cneorum tricoccon L. (Gneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Reproductive ecology of Cneorum tricoccon L. (Cneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Changes in Florets’ Vertical Direction within Inflorescence Affects Pollinator Behavior,and Fitness in <i>Trifolium repens</i>

Ecological interactions between flowers and pollinators greatly affect the reproductive success. To facilitate these interactions, many flowers are known to display their attractive qualities, such as scent emission, flower rewards and floral vertical direction, in a rhythmic fashion. However, less is known about how plants regulate the relationship between these flower traits to adapt to pollinator visiting behavior and increase reproduction success. Here we investigated the adaptive significance of the flower bending from erect to downward in Trifolium repens. We observed the flowering dynamic characteristics (changes of vertical direction of florets, flowering number, pollen grain numbers, pollen viability and stigma receptivity over time after blossom) and the factors affecting the rate of flower bending in T. repens. Then we altered the vertical direction of florets in inflorescence of different types (upright and downward), and compared the pollinator behaviors and female reproductive success. Our results showed that florets opened sequentially in inflorescence, and then bend downwards slowly after flowering. The bending speed of florets was mainly influenced by pollination, and bending angle increased with the prolongation of flowering time, while the pollen germination rate, stigma receptivity and nectar secretion has a rhythm of “low-high-low” during the whole period with the time going. The visiting frequency of all the four species of pollinators on upward flowers was significantly higher than that of downward flowers, and they especially prefer to visit flowers with a bending angle of 30°–60°, when the flowers was exactly of the highest flower rewards (nectar secretion and number of pollen grains), stigma receptivity and pollen germination rate. The seed set ratio and fruit set ratio of upward flowers were significantly higher than downward flowers, but significantly lower than unmanipulated flowers. Our results indicated that the T. repens could increase female and male fitness by accurate pollination. The most suitable flower angle saves pollinators’ visiting energy and enables them to obtain the highest nectar rewards. This coordination between plants and pollinators maximizes the interests of them, which is a crucial factor in initiating specialized plant-pollinator relationships.     

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.     

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.     

Phenology and phenotypic natural selection on the flowering time of a deceit-pollinated tropical orchid, Myrmecophila christinae

BACKGROUND AND AIMS: Flowering phenology is described and the effect of flowering time on pollination success is evaluated in the deceit-pollinated tropical orchid, Myrmecophila christinae. It was expected that, due to this species' deceit pollination strategy and low observed pollinator visit rate, there would be a higher probability of natural selection events favouring individuals flowering away from the population flowering peak. METHODS: The study covers two consecutive years and four populations of M. christinae located along the north coast of the Yucatán Peninsula. For phenological and pollination success data, a total of 110 individuals were monitored weekly in 1998, and 83 individuals in 1999, during all the flowering and fruiting season. KEY RESULTS: The results showed significant differences in the probability of donating and receiving pollen throughout the flowering season. The probability of receiving or donating pollen increased the further an individual flowering was from the flowering peak. Regression analysis showed directional and disruptive phenotypic natural selection gradients, suggesting the presence of selection events unfavourable to flowering during flowering peak, for both male success (pollen removal) and female success (fruit production). However, the intensity and significance of the natural selection events varied between populations from year to year. The variation between seasons and populations was apparently due to variations in the density of reproductive individuals in each population and each season. CONCLUSIONS: As in other deceit-pollinated orchids, natural selection in M. christinae favours individuals flowering early or late in relation to population peak flowering. However, results also suggested a fluctuating regime of selective events act on flowering time of M. christinae.     

How pollinator-mediated mating varies with population size in plants

In most higher plants sexual interactions are mediated by animal pollinators that affect the number and differential reproductive success of mates. The number and sex of breeding individuals in populations are central factors in evolutionary theory, but the quantitative effect of plant population size on pollinator-mediated mating is understudied. We investigated variation in pollen removal (male function) and fruit set (female function) among flowering populations of different size of two bumblebee-and one butterfly-pollinated, rewardless, pollen-limited, hermaphroditic orchid species in Sweden. As the amount of pollen removed from plants by insects (either absolute or proportional) increased, so did the number of pollinations, whereas the proportions of plants with different pollinator-designated functional sex (male, female, hermaphrodite) depended primarily on the ratio between the amount of fruit set and pollen removed within populations. A larger population size was found to have several effects: (1) the total numbers of pollinia removed and fruits set increased; (2) the proportion of pollen removed from plants decreased; (3) the proportion of flowers pollinated decreased in the butterfly-but was not affected in the bumblebee-pollinated species; (4) the ratio between fruits set and pollinia removed increased linearly in the bumblebee-pollinated species but reached a maximum at c. 80 individuals in the butterfly-pollinated species; (5) the numbers of pollinator-designated pure male and hermaphrodite individuals increased; and (6) the variance in pollinium removal, but not fruit set, increased among individuals. These findings empirically verify the basic importance of population size for the mating structure of outcrossing plants, and indicate that selection for female sexual traits is reinforced when population size is smaller while selection for male sexual traits is reinforced when population size is larger.     

Reproductive ecology of the endemic Lecocarpus pinnatifidus (Asteraceae) in an isolated population in the Galápagos Islands

Lecocarpus pinnatifidus is an endemic member of the Asteraceae occurring on only one island in the Galápagos archipelago. The capitula are large with female ray florets and male disc florets. They are self-compatible but this study suggests fruit set is pollen limited. Visits from Xylocopa darwini and other larger insect pollinators are rare, and small insects seem to be the main pollinators. Small insects carry few pollen grains and most likely mediate self-pollinations. Self-compatibility and seed set after selfing are the most common reproductive strategy in the Galápagos Islands and L. pinnatifidus seemingly fits well into this group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 171–180.     

Female Fecundity Is Dependent on Substrate, Rather Than Male Abundance, in the Wind-Pollinated, Dioecious Understory Palm Chamaedorea radicalis

We examined the impact of substrate, population density, and sexual composition on female reproductive success in harvested populations of the tropical understory palm Chamaedorea radicalis . Leaf harvest for the international floral greenery trade does not kill palms, but has been shown to reduce flowering frequency and has been projected to reduce population growth. Because C. radicalis is dioecious and wind-pollinated, a reduction in flowering density from leaf harvest or other anthropogenic factors may lower pollen movement between flowering conspecifics and lower fruit production. Such reduced fecundity in harvested populations can contribute to an Allee effect, where the per capita rate of population growth declines at low density. We tested for these effects by sampling C. radicalis along transects in ten populations, spanning a range of densities, during May (flowering peak) and August (fruiting peak) 2002. We applied path analysis using structural equation modeling (SEM) to test a set of hypothesized relationships between substrate, female size, several population parameters, and female fecundity. SEM models revealed that female fecundity was not dependent on any measure of population density or sexual composition, providing no evidence for an Allee effect. The strongest model ( R ²= 92.9%) related C. radicalis reproductive output only to substrate, female size, flower number, and proportional fruit set. This model revealed that palms on rock outcrops were larger, produced more flowers, and had higher proportional fruit set than palms on the forest floor. This finding suggests that protection of females on outcrops should enhance population growth and viability.     

Maytenus obtusifolia Mart. (Celastraceae): a tropical woody species in a transitional evolutionary stage of the gynodioecy–dioecy pathway

‘Gynodioecy–dioecy’ is one of the pathways by which dioecy can arise from hermaphroditism. Studies on sex determination and development of gynodioecious systems have focused on temperate and/or annual species. Little is known about the evolutionary dynamics of gynodioecy and dioecy in perennial tropical species, where these systems have more frequently evolved. Maytenus obtusifolia is an abundant species in restingas in southeastern Brazil. The sexual system of M. obtusifolia was investigated by studying the floral structure and reproductive biology. We considered the sexual system as an intermediate step in the pathway gynodioecy-dioecy. The characterization of the hermaphrodite morph was complex, because of a gradient of variation in floral morphology and reproductive characteristics (sizes of the style and stigmatic surface, pollen viability, embryo sacs containing hypertrophied synergids, and fruit set). This variation leads to different proportions of functional male and female flowers among hermaphrodite plants and is responsible for the different levels of reproductive success. Female reproductive success and pollen viability were negatively correlated with the hermaphrodite morph (r = ?0.67). The higher fruiting intensity and fruit/flower ratio of females (41 %) compared to hermaphrodites (2 %) and the fact that female plants produce more and better-quality seeds support the female compensation. We suggest that female sterility may be linked to the set of changes in the carpels. The differences in the quality and quantity of pollen grains of hermaphrodite plants, and the similar individual rates of pollen viability observed for three consecutive flowering events, may indicate a relationship with nuclear cytoplasmic sex determination.     

Early blooming's challenges: extended flowering season, diverse pollinator assemblage and the reproductive success of Gynodioecious Daphne laureola

BACKGROUND AND AIMS: The scarcity and unpredictability of active pollinators during late winter in temperate areas tends to favour extended flowering seasons and increased floral longevity in early blooming species, which are usually pollinated by diverse sets of insects. Daphne laureola is a gynodioecious woody perennial that flowers from January to April in southern Spain, a period characterized by cold temperatures, frequent rains and irregular snowfalls. METHODS: Pollinators were excluded at four different times during the flowering season in order to determine the effect of decreased exposure to pollinators on fruit set in female and hermaphrodite individuals. The role of nocturnal and diurnal pollination on reproductive success in each gender was simultaneously evaluated by selective exclusion. KEY RESULTS: A 50 % reduction in the flowering period decreased fruit set of females by 50 %, whereas the corresponding decrease in self-compatible hermaphrodites was only approx. 25 %. Day-active hymenopterans and lepidopterans were infrequent visitors, and nocturnal pollinators were inefficient, suggesting that pollen beetles, Meligethes elongatus, were the main pollinators of D. laureola in the study region. CONCLUSIONS: Beetles were less abundant in pollenless females, although discrimination did not apparently result in pollination limitation of female reproduction. A preference of beetles for sunny locations emphasized the relevance of abiotic conditions for pollination of this early blooming shrub.     

小花风毛菊的性器官在青藏高原的海拔变异

 小花风毛菊(Saussurea parviflora)是菊科风毛菊属的一种多年生草本植物,是该属在青藏高原东缘的一个优势种。该文研究了小花风毛菊11个居群性器官的变异与11个海拔高度(2 100～3 500 m)之间的相关性,以及花丝、花药长和花柱、花柱分枝长与花粉数之间的相关性;探讨了小花风毛菊的性器官是如何随海拔高度发生变异,如何与胁迫环境相适应,实现交配成功,提高繁殖成功率,使其成为该属植物“现代 分布中心"的优势种。结果表明：1)花丝、花药长与海拔高度呈极显著的正相关关系(p<0.01);2)花粉数与海拔高度呈极显著的负相关关系(p<0.01);3)花柱、花柱分枝长及花柱分枝长和雌蕊长的比与海拔高度呈极显著的正相关关系(p<0.01);4)花柱、花柱分枝长和花丝长与花粉数呈显著的负相关关系(p<0.05);5)种子的成熟率与海拔高度呈显著的正相关关系(p<0.05)。说明小花风毛菊种内性器官在特定的生存环境下发生了变异,特别是在花粉数随海拔的升高明显减少以及昆虫的多样性、丰富性和活动性明显降低的情况下,它能够通过花柱分枝长度的变异来提高柱头对传粉昆虫的感受性,能通过种子成熟率的提高来保证高海拔数目显著减少的种子的成功发育,进而在一定程度上导致了该物种繁殖成功率的上升,使它成为青藏高原这一胁迫环境中的优势种。     

喜树开花特性及繁育系统的研究

定位观测了喜树(Camptotheca acuminata Decne)花部构造、开花特性,用杂交指数(OCI)、花粉/胚珠比(P/O)、去雄、套袋、人工授粉等方法测定了喜树的繁育系统.结果表明:喜树花序为由4～6个头状花序组成的聚伞花序,聚伞花序上部的头状花序由两性花组成,下部的花序由雄花组成,两性花雌雄蕊发育正常,雄蕊10枚,雌蕊1枚,下位子房,一室,倒生胚珠;雄花雄蕊发育正常,雌蕊不发育.自然条件下,同一头状花序散粉后2 d,花粉活力最高;两性花在柱头外翻后3～4 d,柱头可受性最强,第6天失去可受性.喜树两性花为半同步雌雄蕊异熟类型,每个头状花序的所有小花同步开放,雄蕊先成熟,表现为雄性时期,雄蕊脱落后,雌蕊成熟,表现为雌性时期,上一级头状花序雌蕊成熟期与下一级头状花序雄蕊成熟期重合.繁育系统检测结果为部分自交和异交,需要传粉者活动才能完成授粉过程.     

Flowering in a bamboo, Melocanna baccifera (Bambusoideae: Poaceae)

Melocanna baccifera (Roxburgh) Kurtz ex Skeels, a species of bamboo introduced to Sri Lanka from India, flowered and set fruit during 2001–2002. Culms that flowered and set fruit died. The incidence of flowering is significant in that flowering took place close to the predicted mast flowering in 2007. At the onset of flowering, inflorescences were predominantly staminate. But later in 2002, bisexual and pistilate flowers also developed leading to fruiting. Both protandry and protogyny were observed in the bisexual florets. Floral characters indicated that the species was mainly out-crossing. Although anther dehiscence released pollen and stigmas were exerted, pollination of stigmas was inefficient. The few stigmas that were naturally pollinated showed limited pollen tube growth. However, fruit set took place. Fruits were very rarely seen to germinate naturally. Many fruits were devoid of an embryo, indicating that parthenocarpy and/or apomixis may have taken place. Excised embryos germinated in vitro .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 287–291.     

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

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

Reproductive allocation and costs in gynodioecious Leucopogon melaleucoides (Ericaceae): implications for the evolution of gender dimorphism

In dioecious species, females typically allocate more resources to reproduction and incur greater costs of reproduction than males. In gynodioecious species, sex-based differences in reproductive allocation (RA) and costs have been less studied. Such knowledge, however, is relevant to address how females establish and increase in frequency in populations. We examine RA and reproductive costs by comparing fruit set, the proportion of biomass allocated to reproduction, and the responses of fruit set and vegetative growth to shoot defoliation in females and hermaphrodites in gynodioecious Leucopogon melaleucoides. Relative to hermaphrodites, females exhibited a two-fold fruit set advantage. Female fruit set increased proportionately with flower number, but hermaphrodite fruit set was reduced on plants with more flowers. Sex-based differences in allocation to other traits were small. Thus, female RA at flowering was similar to hermaphrodite RA, but was 1.4-fold greater at fruiting. Relative to controls, defoliation reduced fruit set and the percentage of shoots that produced new vegetative growth similarly in both sexes. However, females had a lower proportion of shoots with new growth overall. Further, defoliation on females reduced the dry mass of new growth by 44% compared with controls, whereas hermaphrodites were not affected. These results indicate a trade-off between reproduction and vegetative growth, and greater female costs of reproduction, particularly under resource-limiting conditions. In the absence of compensatory traits to offset higher female reproductive costs, such trade-offs have the potential to retard the spread of females in gynodioecious populations.     

Mating patterns and contemporary gene flow by pollen in a large continuous and a small isolated population of the scattered forest tree Sorbus torminalis

The influence of population size and spatial isolation on contemporary gene flow by pollen and mating patterns in temperate forest trees are not well documented, although they are crucial factors in the life history of plant species. We analysed a small, isolated population and a large, continuous population of the insect-pollinated tree species Sorbus torminalis in two consecutive years. The species recently experienced increased habitat fragmentation due to altered forest management leading to forests with closed canopies. We estimated individual plant size, percentage of flowering trees, intensity of flowering, degree of fruiting and seed set per fruit, and we determined mating patterns, pollen flow distances and external gene flow in a genetic paternity analysis based on microsatellite markers. We found clear effects of small population size and spatial isolation in S. torminalis. Compared with the large, continuous population, the small and isolated population harboured a lower percentage of flowering trees, showed less intense flowering, lower fruiting, less developed seeds per fruit, increased selfing and received less immigrant pollen. However, the negative inbreeding coefficients (F(IS)) of offspring indicated that this did not result in inbred seed at the population level. We also show that flowering, fruiting and pollen flow patterns varied among years, the latter being affected by the size of individuals. Though our study was unreplicated at the factor level (i.e. isolated vs non-isolated populations), it shows that small and spatially isolated populations of S. torminalis may also be genetically isolated, but that their progeny is not necessarily more inbred.     

A multi-year study of factors affecting fruit production in Aristolochia paucinervis (Aristolochiaceae)

Pollen limitation, resource limitation, fruit abortion, and predation have all been proposed as factors explaining low fruit set in hermaphroditic plants. We conducted a 5-year study combining field observations and pollination experiments to determine the causes of the low fruit set in Aristolochia paucinervis, a Mediterranean species with a specialized pollination system in two populations in SW Spain. Fruit initiation was markedly low, and between 28.6 and 75.0% of the flowering stems did not initiate any fruit. In most flowers, the number of germinated pollen grains was less than the number of ovules, and supplemental pollination significantly increased fruiting, indicating deficient pollination. In A. paucinervis, autonomous self-pollination seems to be a decisive factor in fruit production because the number of germinated pollen and the fruit set from flowers bagged before anthesis were similar to those in free-pollinated flowers. Only in 2005 did flowers that were successfully pollinated outnumber ripened fruits, suggesting that other factors limit fruiting. We found a significant positive correlation between tuber mass and fruit set. Deficient pollination and lack of resources could explain the low fruit set, but the relative consequences seem to vary spatially and temporarily.