Ovule pre-emption and pollen limitation in a self-fertile perennial herb (Blandfordia grandiflora,Liliaceae)

The extent, frequency and causes of pollenlimited seed production were examined in partially selffertile populations of Blandfordia grandiflora for 2 years. Percentage seed set of open-pollinated plants (50–57%) did not differ within or between years, and was about 19% less than experimentally cross-pollinated plants (70–75%). Floral visits by honeybees did not differ through the flowering season and the number of pollen grains deposited on stigmas within 12 h of flowers opening exceeded the number of ovules per flower, indicating that the quality rather than the quantity of pollination limited seed set. Pollen limitation was caused by concurrent self- and cross-pollination and the subsequent abortion of some selfed ovules due to inbreeding depression. Natural seed set (55%) was intermediate between selfed (43%) and crossed (75%) flowers and was not increased when flowers that had been available to pollinators for 24 h were hand cross-pollinated, suggesting that ovules were already fertilized. Similarly, experimental pollination with both cross and self pollen within 24 h of flowers opening did not increase seed set relative to natural seed set, indicating that both cross- and self-fertilizations had occurred. In contrast, when selfing followed crossing by 48 h, or vice versa, seed set did not differ from crossed-only or selfed-only flowers, respectively, indicating that ovules were pre-empted by the first pollination. Collectively, these results indicate that under natural conditions self pollen pre-empts ovules, rendering them unavailable for cross-fertilization. This selfing reduces fecundity by 50%, as estimated from the natural production of cross seeds when selfing was prevented. Consequently, selection should favour floral traits, such as increased stigma-anther separation or protandry, that reduce interference between male and female functions that leads to selfing.     

Many to flower, few to fruit: the reproductive biology of Hamamelis virginiana (Hamamelidaceae)

Hamamelis virginiana flowers from late September to late November. In 1977, we began studying the reproductive biology of this eastern North American arborescent shrub by examining floral phenology and rewards, pollen-ovule ratios, breeding system, pollination, pollinator and resource limitation, and seed dispersal. The homogamous, self-incompatible flowers emit a faint odor, bear nectar with sucrose ratios typical of bee- and fly-pollinated flowers, and produce abundant sticky pollen. Flowers were visited infrequently by insects representing six orders. Flies were the most common floral visitors, specifically members of the genus Bradysia, but small bees also carried high percentages of Hamamelis pollen. Despite high pollen/ovule ratios (11?445 grains/ovule), bees and flies are likely pollinators, as experiments indicate wind pollination is less likely. Pollen quantity and resource availability did not appear to limit reproductive output, but pollen quality did. Tests of >40?000 flowers showed natural fruit set to be <1%. The flowering time, breeding system, and clumped distribution of plants, likely due in part to limited seed dispersal, combine to yield this remarkably low fruit set. Because all other species of Hamamelis flower from late winter to early summer, it may be that H. virginiana evolved a fall flowering phenology to avoid competition for pollinators with the closely related H. vernalis.     

Testing hypotheses for excess flower production and low fruit-to-flower ratios in a pollinating seed-consuming mutualism

Pollinator attraction, pollen limitation, resource limitation, pollen donation and selective fruit abortion have all been proposed as processes explaining why hermaphroditic plants commonly produce many more flowers than mature fruit. We conducted a series of experiments in Arizona to investigate low fruit-to-flower ratios in senita cacti, which rely exclusively on pollinating seed-consumers. Selective abortion of fruit based on seed predators is of particular interest in this case because plants relying on pollinating seed-consumers are predicted to have such a mechanism to minimize seed loss. Pollinator attraction and pollen dispersal increased with flower number, but fruit set did not, refuting the hypothesis that excess flowers increase fruit set by attracting more pollinators. Fruit set of natural- and hand-pollinated flowers were not different, supporting the resource, rather than pollen, limitation hypothesis. Senita did abort fruit, but not selectively based on pollen quantity, pollen donors, or seed predators. Collectively, these results are consistent with sex allocation theory in that resource allocation to excess flower production can increase pollen dispersal and the male fitness function of flowers, but consequently results in reduced resources available for fruit set. Inconsistent with sex allocation theory, however, fruit production and the female fitness function of flowers may actually increase with flower production. This is because excess flower production lowers pollinator-to-flower ratios and results in fruit abortion, both of which limit the abundance and hence oviposition rates, of pre-dispersal seed predators.     

The interplay of pollination,costs of reproduction and plant size in maternal fertility limitation in perennial <Emphasis Type="Italic">Paeonia officinalis</Emphasis>

Although several factors can limit female fertility in perennial plants, rarely have they been jointly studied in a single species over several years. In this study we experimentally manipulate seed production and simultaneously analyse the potential contribution of pollen limitation, costs of reproduction and plant size to variation in seed output over a 3-year period in the perennial herb Paeonia officinalis, in southern France. Since this rare species is threatened by forest closure in many sites we also examine the causes of female fertility variation in relation to habitat closure (open habitat vs. woodland). P. officinalis has a partial self-incompatibility system and only very low ability for autonomous self pollination in the absence of pollinators. However, supplementary pollination of individual plants in three consecutive years did not significantly increase seed production above natural levels. Forest closure was associated with a decline in ovule and seed production, which again was not due to pollen limitation since supplementary pollination had no significant effect on seed set in the woodland habitat. Comparison of the maternal fertility of plants which were previously excluded from reproduction with those which were hand pollinated to maximise seed set in two previous years produced no evidence that seed production in year three is limited by costs associated with prior reproduction. Likewise, flowering probability was not related to prior seed production but was however positively related to plant size. The absence of any influence of pollen limitation or prior reproduction on seed production suggests that sub-maximal seed production in long-lived perennial herbs may be part of a size-dependent strategy that maximises life-time seed production and fitness without compromising survival.     

Seasonal changes in pollen limitation and femaleness along the snowmelt gradient in a distylous alpine herb,Primula modesta

Flowering phenology of alpine plants is strongly determined by the timing of snowmelt, and the conditions of pollination of widely distributed plants vary greatly during their flowering season. We examined the reproductive success of the distylous alpine herb, Primula modesta, along the snowmelt gradient under natural conditions, and compared it with the result of artificial pollination experiments. In addition, the compositions and visit frequencies of pollinators to the flower of P. modesta were examined during the flowering period. The pin and thrum plants of P. modesta growing at the same site have an equal ability to produce seeds if a sufficient amount of legitimate pollen grains are deposited on the stigma surface. However, under natural conditions, their seed‐set success was often (even if not always) restricted by pollen limitation, and the functional gender of the pin and thrum plants biased to the female and male, respectively, associated with their growing sites. These variations were not ascribed to resource limitation nor biased morph ratio but to the seasonal changes in pollination situations, a replacement of pollinator types from long‐ to short‐tongued pollinators resulted in unidirectional pollen transfer from long stamens (thrum plants) to long styles (pin plants). The functional gender specialization may enhance the evolution of dioecy from heterostyly, but the severe pollen limitation may cause the breakdown of heterostyly into homostyly. To consider the evolutionary pathway of heterostylous plants, an accumulation of the empirical data is required demonstrating how phenological synchrony between plants and pollinators is decided and to what degree this relationship is stable over years, along with estimates of selection and gene flow in individual plants.     

Patterns of pollen quantity and quality limitation of pre‐zygotic reproduction in Mimulus guttatus vary with co‐flowering community context

Pollen quantity limitation has been widely recognized as one of the main causes of plant reproductive failure in nature. However, its negative effects on fruit and seed production have been often confounded with those of low pollen quality (i.e. low conspecific pollen viability and/or slow pollen tube growth rate). The lack of differentiation between these two aspects of pollen limitation has resulted not only in a potential overestimation of the incidence of pollen quantity limitation but has led to a poor understanding of the factors and mechanisms that affect pollen quality limitation. Knowledge of the relative importance and underlying causes of both aspects of pollen limitation (quantity and quality) is required to fully understand the ecological and evolutionary consequences of pollen limitation in natural populations. Co‐flowering community context (e.g. species richness and conspecific density), in particular, can be an important driver of overall pollen limitation. However, how pollen quantity and quality limitation vary separately with the community context has not previously been examined, even though they may vary differently as they arise from different mechanisms, e.g. insect visitation rate versus inbreeding depression. Here we evaluate the effect of co‐flowering diversity and conspecific density on the relative importance of pollen quantity and quality limitation for Mimulus guttatus pre‐zygotic reproduction (i.e. pollen tube success) at serpentine seeps in California over two years. We found overall pollen limitation of pre‐zygotic reproductive success at all seeps regardless of the co‐flowering context. However, plants in high‐diversity/low‐ conspecific density communities were mostly limited by pollen quantity, whereas plants in low‐diversity/high‐conspecific density ones experienced stronger quality limitation – a pattern that was consistent across years. These results not only highlight the importance of conducting comprehensive studies on pollen limitation that evaluate both, its quantity and quality aspects, but are the first to show how their relative contribution to overall pollen limitation can vary with extrinsic factors such as the co‐flowering community context.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Effects of pollen quality and quantity on pollen limitation in Crataegus monogyna (Rosaceae) in NW Spain

Pollen limitation occurs when plants produce less fruits and/or seeds than they would with adequate pollen receipt. If the addition of cross-pollen to stigmas increases fruit/seed production, it is interpreted as an evidence of pollen limitation. Much of the limitation may be associated with the quality rather than quantity of pollen; however, most studies do not discriminate between the two, which may lead to misinterpretation of the results. We studied the effects of quality and quantity of pollen on the reproduction of a northern Spanish population of Crataegus monogyna. The treatments included self- and cross-pollination, and supplementation to open and bagged flowers. The response variables considered were number of pollen grains per stigma, pollen tubes per style, and initial and final fruit set. In the Cantabrian range, C. monogyna requires insect pollinators to set fruit and is partially self-incompatible. We found that the number of pollen tubes did not differ between cross- and self-pollination treatments; however, self-pollinated flowers set less fruits than flowers that received pure cross-pollen or were supplemented with both cross- and self-pollen. The experimental design allowed us to infer qualitative rather than quantitative pollen limitation. Comparison of the number of pollen grains and tubes, and initial and final fruit set among pollination treatments suggested post-zygotic embryo selection against selfed progeny.     

Pollen quality limits seed set in Burchardia umbellata (Colchicaceae)

In self-incompatible plants, interference by self pollen or genetically related pollen can potentially exacerbate pollen limitation, although this has rarely been demonstrated. We examined the breeding system, pollen limitation, and pollen interference using self- and cross- pollinations and pollen supplementations in Burchardia umbellata, an insect-pollinated lilioid monocot. Ovule fertilization and seed set were less following selfing than crossing (22 vs. 78% and 4 vs. 73%, respectively), indicating partial self-incompatibility. Flowers were partially protandrous, and flowers opened concurrently on plants potentially allowing self pollen interference. Natural seed set was pollen limited and varied within and among years, probably due to variation in flowering plant density. Interference by self or genetically related pollen caused pollen limitation as evidenced by increased seed set of bagged cross-pollinated plants compared to unbagged pollen-supplemented plants in two years. In 1996, both fertilization and seed set increased in response to cross-pollination, indicating that interference occurred in the style and ovary. In 1997, only seed set increased after cross-pollination indicating that interference occurred in the ovary. Inappropriate pollen deposition may contribute to pollen limitation more often than previously recognized and should select for floral traits that decrease deposition of self or related pollen.     

Variability in Pollen Load: Implications for Reproduction and Seedling Vigor in a Rare Plant, Silene douglasii var. Oraria

Habitat fragmentation can markedly influence the levels of pollen deposition and seed production in natural populations, and rare plants may be especially susceptible to any associated reductions in pollen quantity and quality. In order to ascertain the potential for pollen limitation of maternal fitness in a rare plant, Silene douglasii var. oraria, which is endemic to western coastal prairies, we counted ovules and measured conspecific and heterospecific pollen deposition on stigmas collected from open-pollinated plants. We further investigated the effect of increasing pollen intensity on fruit production, seed number and weight, as well as several measures of progeny vigor. Three levels of outcross pollen were added to plant stigmas for comparison with autogamous and open pollination in the largest naturally occurring population. Both seed and fruit production were significantly greater (P<0.05) for supplemented versus nonsupplemented stigmas, but flowers receiving different levels of pollen addition were statistically indistinguishable. Seed germination and seedling survival were also lowest for the offspring of nonsupplemented flowers; however, in natural populations, opportunities for pollen competition are very limited since open-pollinated flowers averaged fewer viable pollen grains than ovules. Seed production was equivalent for open- and autogamously pollinated flowers in 1996, indicating that natural pollen transfer may have involved mostly self pollen. Overall, the low reproductive success of var. oraria likely reflects both low pollen quantity and quality. Multiyear empirical studies of pollen intensity in field populations are needed so that we can better understand the fitness consequences of pollen limitation in rare perennials.     

Pollination in Verbascum thapsus (Scrophulariaceae): the advantage of being tall

According to the "effective pollination" hypothesis, tall stature resulting from strong apical dominance attracts greater pollinator visitation, thus allowing larger pollen loads and/or greater outcrossing rates, which in turn produces more vigorous offspring with greater genotypic variability and/or less inbreeding depression. Components of this hypothesis were tested in Verbascum thapsus, which commonly grows unbranched to over 2 m tall with strong apical dominance suppressing all axillary meristems. A natural population survey indicated that plants with visiting pollinators were significantly taller than their nearest neighboring individuals not possessing a visiting pollinator. Plants in natural populations with excluded pollinators produced seeds via a delayed selfing mechanism. However, delayed selfing under pollinator exclusion resulted in only 75% of the seed set obtained with natural pollinators. Under natural pollination, emasculated flowers experienced a 50% reduction in pollen deposition by the time of flower closure but only a 5% reduction in seed set relative to intact flowers. Hence, taller plants attracted more pollinators and maximum seed set could not be achieved without pollinators. Comparison of seed set and seed mass in plants that were artificially selfed and artificially crossed (in both the greenhouse and in natural populations) indicated that plants were fully self-compatible with no evidence of early-acting inbreeding depression. However, this does not exclude the possibility that inbreeding depression is manifested in later life stages. The results suggest that V. thapsus has a mixed mating system with potential for reproductive assurance and various levels of outcrossing depending on variables affecting pollinator availability (e.g., population size).     

Does self-pollination provide reproductive Assurance in Aquilegia Canadensis (Ranunculaceae)?

The ability to produce seeds when pollinators or potential mates are scarce is thought to be one of the main advantages of self-fertilization in flowering plants. However, whether autonomous selfing increases seed set in natural populations has seldom been tested, and even fewer studies have evaluated the advantage of selfing across a gradient of pollen availability. This study examines the fertility consequences of autonomous selfing in Aquilegia canadensis (Ranunculaceae), a short-lived, spring-flowering perennial typically found in small, patchy populations on rock outcrops. We used a pollinator exclusion experiment to confirm reports that A. canadensis has a well-developed capacity for autonomous selfing resulting from incomplete protogyny and close proximity of stigmas and anthers during dehiscence. Flowers excluded from pollinators set 87% as many seeds per carpel (X +/- 1 SE = 7.1 +/- 1.4 seeds) as hand-pollinated flowers (8.1 +/- 1.3 seeds), and seed production in unpollinated flowers correlated negatively with the distance between stigmas and anthers (r = -0.46). Autonomous selfing could be potentially valuable in providing reproductive assurance because only 2.7 +/- 0.5 pollen grains were deposited on each stigma before anther dehiscence, compared to 134.1 +/- 17.9 pollen grains by the end of anther dehiscence. However, prevention of autonomous selfing by anther removal before dehiscence did not decrease seed set, even for plants at low plant densities where outcross pollen may have been in short supply. Emasculated flowers set as many seeds per carpel (9.3 +/- 0.9) as intact flowers (8.4 +/- 1.1), indicating that sufficient cross pollen is deposited for full seed set. These results do not support the hypothesis that autonomous selfing by A. canadensis has been selected because it provides reproductive assurance.     

Biocontrol attack increases pollen limitation under some circumstances in the invasive plant Centaurea solstitialis

Herbivore damage often deters pollinator visitation and many invasive plants in North America are pollinator-dependent. This has important implications for the biological control of invasive plants because it means that agents that deter pollinators may have a larger than expected impact on the plant. Yet interactions between pollinators and biocontrol agents are rarely evaluated. Centaurea solstitialis, one of the most problematic invasive species in California, is dependent on pollinators for reproduction. I factorially manipulated infection by a biocontrol pathogen and pollen supplementation to test for (1) pollen limitation in C. solstitialis, (2) whether infection increased pollen limitation, and (3) whether this varied across a soil moisture gradient. Plants growing on north-facing slopes where soil moisture was higher experienced mild pollen limitation in the absence of the pathogen and more pronounced pollen limitation when they were infected. Plants on drier south-facing slopes did not suffer from pollen limitation but instead appeared to suffer from resource limitation. Pathogen infection directly reduced seed set in C. solstitialis by 67–72 %. On north-facing slopes, infection had an additional, indirect effect by increasing the degree of pollen limitation plants experienced. The trait that mediates this indirect pathogen–pollinator interaction is the number of inflorescences plants produced: infected plants made fewer inflorescences which led to greater pollen limitation. Although in the present study this outcome is dependent on abiotic factors that vary over small spatial scales, exploiting other invasive plants’ dependence on pollinators by selecting agents that deter visitation may enhance agent impact.     

Reproductive consequences of mate quantity versus mate diversity in a wind-pollinated plant

Since most pollen travels limited distances in wind-pollinated plants, both the local quantity and diversity of mates may limit female reproductive success. Yet little evidence exists on their relative contribution, despite the importance of viable seed production to population dynamics.To study how variation in female reproductive success is affected by the quantity versus the diversity of surrounding mates contributing pollen, we integrated pollination experiments, data on natural seed set and seed viability, and AFLP genetic marker data in the wind-pollinated dioecious clonal forest herb Mercurialis perennis.Pollination experiments indicated weak quantitative pollen limitation effects on seed set. Among-population crosses showed reduced seed viability, suggesting outbreeding depression due to genetic divergence. Pollination with pollen from a single source did not negatively affect reproductive success. These findings were consistent with results of the survey of natural female reproductive success. Seed set decreased with the distance to males in a female plants’ local neighborhood, suggesting a shortage of pollen in isolated female plants, and increased with the degree of local genetic diversity. Spatial isolation to other populations and population size did not affect seed set. None of these variables were related to seed viability.We conclude that pollen movement in M. perennis is likely very limited. Both male proximity and the local degree of genetic diversity influenced female reproductive success.     

Limits to reproductive success of Sarracenia purpurea (Sarraceniaceae)

Plant biologists have an enduring interest in assessing components of plant fitness and determining limits to seed set. Consequently, the relative contributions of resource and pollinator availability have been documented for a large number of plant species. We experimentally examined the roles of resource and pollen availability on seed set by the northern pitcher plant Sarracenia purpurea. We were able to distinguish the relative contributions of carbon (photosynthate) and mineral nutrients (nitrogen) to reproductive success. We also determined potential pollinators of this species. The bees Bombus affinis and Augochlorella aurata and the fly Fletcherimyia fletcheri were the only floral visitors to S. purpurea that collected pollen. Supplemental pollination increased seed set by <10%, a much lower percentage than would be expected, given data from noncarnivorous, animal-pollinated taxa. Seed set was reduced by 14% in plants that could not capture prey and by another 23% in plants whose pitcher-shaped leaves were also prevented from photosynthesizing. We conclude that resources are more important than pollen availability in determining seed set by this pitcher plant and that reproductive output may be another "cost" of the carnivorous habit.     

Effects of floral herbivory on foraging behaviour of bumblebees and female reproductive success in Pedicularis gruina (Orobanchaceae)

The effects of floral herbivores on floral traits may result in alterations in pollinator foraging behaviour and subsequently influence plant reproductive success. Fed-upon plants may have evolved mechanisms to compensate for herbivore-related decreased fecundity. We conducted a series of field experiments to determine the relative contribution of floral herbivores and pollinators to female reproductive success in an alpine herb, Pedicularis gruina, in two natural populations over two consecutive years. Experimental manipulations included bagging, hand supplemental, geitonogamous pollination, and simulated floral herbivory. Bumblebees not only avoided damaged flowers and plants but also decreased successive visits of flowers in damaged plants, and the latter may reduce the level of geitonogamy. Although seed set per fruit within damaged plants was higher than that in intact plants, total seed number in damaged plants was less than that in intact plants, since floral herbivory-mediated pollinator limitation led to a sharp reduction of fruit set. Overall, the results suggest that resource reallocation within inflorescences of damaged plants may partially compensate for a reduction in seed production. Additionally, a novel finding was the decrease in successive within-plant bumblebee visits following floral herbivory. This may increase seed quantity and quality of P. gruina since self-compatible species exhibit inbreeding depression. The patterns of compensation of herbivory and its consequences reported in this study give an insight into the combined effects of interactions between floral herbivory and pollination on plant reproductive fitness.     

Effects of pollen shortage and self-pollination on seed production of an endangered tree, Magnolia stellata

BACKGROUND AND AIMS: Pollen limitation is a significant determinant of seed production, and can result from both insufficient pollen quantity (pollen shortage) and quality (mainly relating to self-pollination). For animal-pollinated tree species with large floral displays, pollen limitation may be determined by a balance between increased pollen quantity due to increased attractiveness for pollinators, countered by increased self-pollination due to increased geitonogamy. The contributions of pollen shortage and self-pollination on seed production were quantitatively examined in the natural pollination of an insect-pollinated, dichogamous, endangered tree, Magnolia stellata, which has a large, showy floral display. METHODS: Manual self- and cross-pollinations were conducted to determine the effects of selfing on seed production. The outcrossing rate was measured using microsatellite analyses of open-pollinated seeds, and the embryo mortality rate caused by self-pollination was indirectly estimated. The frequency of ovule mortality due to pollen shortage was also inferred using the embryo mortality and ovule survival rates from natural pollination. KEY RESULTS: The average fruit set, seed set per fruit, and ovule survival rate per tree from hand cross-pollination were 1.37, 3.15, and 3.34 times higher than those from hand self-pollination, respectively, indicating that self-pollination causes inbreeding depression for fruit and seed set. The multilocus-outcrossing rate (t(m)) was intermediate, 0.632, and the primary selfing rate was 0.657. This indicates that frequent geitonogamous selfing occurs. The ovule mortality rate due to pollen shortage and the embryo mortality rate due to self-pollination were estimated to be 80.8 % and 45.9 %, respectively. CONCLUSIONS: It is concluded that seed production of M. stellata is strongly limited by both pollen shortage and self-pollination. Inefficient beetle-pollination and the automimicry system via asynchronous flowering might be responsible for the high level of pollen shortage and frequent geitonogamy. This is despite a large, showy floral display and the dichogamous system of the species.     

Proximity to forest edge does not affect crop production despite pollen limitation

A decline in pollination function has been linked to agriculture expansion and intensification. In northwest Argentina, pollinator visits to grapefruit, a self-compatible but pollinator-dependent crop, decline by approximately 50% at 1km from forest edges. We evaluated whether this decrease in visitation also reduces the pollination service in this crop. We analysed the quantity and quality of pollen deposited on stigmas, and associated limitation of fruit production at increasing distances (edge: 10, 100, 500 and 1000m) from the remnants of Yungas forest. We also examined the quantitative and qualitative efficiency of honeybees as pollen vectors. Pollen receipt and pollen tubes in styles decreased with increasing distance from forest edge; however, this decline did not affect fruit production. Supplementation of natural pollen with self- and cross-pollen revealed that both pollen quantity and quality limited fruit production. Despite pollen limitation, honeybees cannot raise fruit production because they often do not deposit sufficient high-quality pollen per visit to elicit fruit development. However, declines in visitation frequency well below seven visits during a flower's lifespan could decrease production beyond current yields. In this context, the preservation of forest remnants, which act as pollinator sources, could contribute to resilience in crop production. Like wild plants, pollen limitation of the yield among animal-pollinated crops may be common and indicative not only of pollinator scarcity, but also of poor pollination quality, whereby pollinator efficiency, rather than just abundance, can play a broader role than previously appreciated.     

Testing for ecological and genetic Allee effects in the invasive shrub Senna didymobotrya (Fabaceae)

For an introduced plant species to become invasive, it must be able to reproduce even in initially small populations. We tested for Allee effects (reduced reproductive performance of individuals in small populations) in the nonclonal, buzz-pollinated shrub Senna didymobotrya in its invasive range in South Africa. The species is self-compatible, but we found that in its invasive range in South Africa it requires pollinators to set seed. Nearly all stigmas (90%) received pollen, but natural fruit set was very low (3-20%). Pollen receipt and fruit set were not significantly correlated with population size. We thus found no evidence for an ecological Allee effect arising from pollen limitation in small populations. Offspring seedling performance, measured in terms of stem volume and leaf area, was also not significantly correlated with the number of plants in the source population, indicating that genetic Allee effects, such as inbreeding depression, are either absent or of such a small magnitude that they would be unlikely to limit further spread of S. didymobotrya in South Africa.     

Variation of pollination and resource limitation in a low seed-set tree, Liriodendron chinense (Magnoliaceae)

Pollen limitation and resource limitation have been documented as the major factors responsible for plants commonly producing more ovules than seeds, but few studies have examined pollen deposition directly in natural populations at different sites and times. We investigated the causes of low seed set in four populations of Liriodendron chinense (Magnoliaceae), an insect‐pollinated endangered tree endemic to southern China, over 2–3 years. One pistil potentially produces two ovules. The number of pistils per flower varies among populations, but in three of the four populations the variation in a given population was not significantly different among years. Overall, populations with higher pistil numbers tend to set more seeds per flower, but a positive correlation between pistil numbers and seed production per flower was observed in only one of the four populations. The numbers of pollen grains deposited per stigma varied from 0 to 60. The proportion of pollinated stigmas per flower ranged from 44% to 88% among populations and years. The numbers of pollen grains deposited per stigma and the percentages of pollinated stigmas were significantly different between populations, and two populations showed significant differences between years. A positive correlation between stigmatic pollen load and seed set was sought in ten population‐by‐year combinations but, in a given population, high stigmatic pollen loads did not always result in high seed set. Examination of pollen deposition, pistil and seed production over several sites and years showed that in addition to pollination, other factors such as resource or genetic loads were likely to limit the (lower than 10%) seed set in L. chinense. It appears that small, isolated populations experience severe pollination limitation; one population studied had seed/ovule ratios of 0.84% and 1.88% in 1995 and 1996. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140 , 31–38.