Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

Floral biology and pollination mechanisms of four Mexico-endemic Fuchsia species with contrasting reproductive systems

Aims The evolution of the outstanding variation of reproductive systems in angiosperms has been considered an important driver of lineage diversification. Closely related hermaphroditic and dioecious species with biotic pollination provide the opportunity to study and compare traits related to pollinator attraction and their consequences on reproductive components. A higher predictability of pollination syndromes is expected in dioecious species, which are dependent on pollinators, than in self-compatible hermaphroditic taxa. Dioecious species may suffer pollen limitation depending on the quality of floral rewards and the kind and abundance of pollinators, whereas no pollen limitation is expected in hermaphroditic species with autonomous self-pollination. Additionally, in the absence of pollen limitation, more or better seeds are expected in dioecious species, according to the sexual specialization hypothesis.Methods In natural populations of the hermaphroditic Fuchsia fulgens and Fuchsia arborescens and dioecious Fuchsia parviflora and Fuchsia obconica, all endemic to Mexico, we first described flower phenology, flower production and longevity and nectar volume and concentration. Then, we evaluated the correspondence between floral visitors and pollination syndromes. In hermaphrodite plants, we determined the level of herkogamy and the potential for autonomous self-pollination. Finally, we evaluated the effect of pollen limitation on fruit set and seed number and assessed seed germination for all species.Important findings In contrast to our prediction, dioecious species did not show a higher correspondence between pollination syndromes and floral visitors than did hermaphrodites; however, male flowers exhibited a higher correspondence than female flowers. No pollen limitation was detected in dioecious species, for which visitation rate did not differ between male and female flowers. The hermaphroditic F. fulgens showed pollen limitation for seed number, despite the presence of autonomous selfing. Fruit set from autonomous pollination was higher in F. arborescens, which showed a lower level of herkogamy compared with F. fulgens. Finally, dioecious species produced fewer but heavier seeds compared with hermaphrodite species. Although Fuchsia is classified as an outcrossing genus, both hermaphroditic species showed autonomous self-pollination. The heavier but lower number of seeds per fruit in dioecious species may be related to the more efficient resource allocation expected from sexual specialization. This could play an important role in the evolution of dioecy; however, a comparative phylogenetic approach is required to confirm this hypothesis.     

ECOLOGICAL CORRELATES OF REPRODUCTIVE TRAITS OF MEXICAN RAIN FOREST TREES

Sexual systems of 139 tree species from a tropical rain forest at Los Tuxtlas, Mexico were investigated to: 1) estimate the relative proportions of hermaphroditic, monoecious, and dioecious species; 2) describe flowers, fruits, and seeds in terms of size and weight; 3) describe flowering and fruiting phenology; and 4) correlate sexuality to pollination and dispersal syndromes, and the successional status occupied in the forest. Hermaphroditism occurred in 63% of the species, monoecism in 9%, and dioecy in 27%. Nondioecious species had larger flowers, but dioecious species had more seeds per fruit. The dioecious condition was associated with small flowers pollinated with unspecialized insects and fleshy fruits dispersed by animals at both species and generic levels. Reproductive traits were more correlated among nondioecious species than dioecious species. Pioneer species had more seeds per fruit, and longer flowering and fruiting periods, but persistent species produced heavier seeds and fruits. Flower and fruit morphological traits, sexual systems, and tree guilds are related in a comprehensive way, and a flow model based on data from this study is presented.     

雄全异株植物瘿椒树（省沽油科）的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树（Tapiscia sinensis Oliv.）的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科（Apidae）和食蚜蝇科（Syrphidae）昆虫,访花高峰期为8：30-10：30。维持瘿椒树雄全异株的可能机制是：雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

雄全异株植物瘿椒树(省沽油科)的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树(Tapiscia sinensis Oliv.)的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科(Apidae)和食蚜蝇科(Syrphidae)昆虫,访花高峰期为8:30-10:30。维持瘿椒树雄全异株的可能机制是:雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

Roots, shoots and reproduction: sexual dimorphism in size and costs of reproductive allocation in an annual herb

Females tend to be smaller than males in woody dioecious plant species, but they tend to be larger in herbs. The smaller size of females in woody species has been attributed to higher reproductive costs, yet no satisfactory explanation has been provided for their larger size in herbs. Because herbs have higher nitrogen concentrations in their tissues than woody plants, and because pollen is particularly rich in nitrogen, we predicted that male growth would be more compromised by reproduction than female growth. To test this hypothesis, we conducted three experiments on the annual dioecious herb Mercurialis annua. First, we compared the timing of reproduction between males and females and found that males started flowering earlier than females; early flowering is expected to compromise growth more than later flowering. Second, we compared plants allowed to flower with those prevented from flowering by experimental debudding and found that males incurred a higher reproductive cost than females in terms of both biomass and, particularly, nitrogen. Third, we grew plants under varying levels of nitrogen availability and found that although sexual size dimorphism was unaffected by nitrogen, females, but not males, decreased their relative allocation to both roots and reproduction under high nitrogen availability. We propose that males deal with the high cost of pollen production in terms of nitrogen by allocating biomass to nitrogen-harvesting roots, whereas females pay for carbon-rich seeds and fruits by investing in photosynthetic organs. Sexual dimorphism would thus seem to be the outcome of allocation to above- versus below-ground sinks that supply resources (carbon versus nitrogen) limiting the female and male reproduction differentially.     

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。     

Pollen-Ovule Ratio and Gamete Investment in Pedicularis （Orobanchaceae）

The Pedicularis species provides ideal materials to study floral evolution because of their substantial flower variation based on a narrow genetic basis, even though they are almost exclusively pollinated by bumblebee. These traits allow us to detect the evolutionary trends of floral parameters without considering genetic background and the difference of pollination vectors. The pollen-ovule ratio is widely used to estimate the pattern of resource investment in two sexual functions in flowering plants. Forty species representing all of the corolla types in Pedicularls were used to study pollen-ovule ratio, gamete investment, and their correlations. Results show that pollen-ovule ratio does not differ among both different corolla types and taxonomic groups. It is therefore suggested that pollen-ovule ratio should be a parallel evolution. The correlations between pollen-ovule ratio and pollen size （-）, and ovule size （＋） can be successfully explained in terms of sex allocation theory. The biological significance of such relationships was also discussed. Additionally, we analyzed the pattern of resource investment into female gamete, which has been somewhat neglected, and found that plants have different patterns of gamete investment between the two sexual functions.     

Reproductive uncertainty and the relative competitiveness of simultaneous hermaphroditism versus dioecy

Hermaphroditism is typically associated with a sedentary existence, whereas dioecy is associated with mobility. This pattern is reflected within flowering plants, as dioecious species commonly possess traits that promote high dispersal. We investigated these associations with three population dynamics models (an individual-based simulation and two mathematical models, one deterministic and the other stochastic) that allowed us to examine competition for space between a hermaphroditic and dioecious species from different perspectives. The competing species are identical in every way but their sexual system. Separation of the sexes increases the variances of pollen import and seed dispersal for the dioecious species. These variances propagate through subsequent reproductive processes and ultimately reduce mean recruitment as a result of nonlinear averaging (Jensen's inequality). A dioecious species could overcome this disadvantage simply by producing more gametes than hermaphrodites; however, in line with the association with mobility, selection on dioecious species should also favor traits that reduce reproductive uncertainty, such as extensive dispersal.     

Pollination biology of the Proteaceae in Australia and southern Africa

Proteaceae are most diverse in southern Africa and Australia, especially in the south-western portions of these regions. Most genera have some species in flower at all times of the year, although generally there is a preponderance of species that flower between late winter and early summer. The one genus that is an exception to this generalization is Banksia, which either has approximately the same percentage of species in flower at various times of the year (southwestern Australia) or peaks in autumn (southeastern Australia). Within particular communities, opportunities for hybridization among congeneric species are minimized by staggered flowering times, different pollen vectors and/or various incompatibility mechanisms. Birds, mammals and arthropods have been identified as visitors to the inflorescences of many Proteaceae. The most common avian visitors to the majority of genera in Australia are honeyeaters, although lorikeets, silvereyes and approximately 40 other species sometimes may be important. Sugarbirds and sunbirds are seen most frequently at inflorescences of Protea, Leucospermum and Mimetes in southern Africa, although they rarely visit other genera. In most cases, avian visitors forage in a manner that permits the acquisition and transfer of pollen. Limited evidence supports the hypothesis that birds are selective in their choice of inflorescences, responding to morphological and/or colour changes and usually visiting those inflorescences that offer the greatest nectar rewards. Arthropods may be equally selective, although it is possible that only the larger moths, bees and beetles are important pollinators, even for those plant species that rely entirely on arthropods for pollen transfer. Mammals are pollen vectors for some Proteaceae, especially those that have geoflorous and/or cryptic inflorescences. In Australia, small marsupials may be the most important mammalian pollinators, although rodents fill this niche in at least some southern African habitats. All but two genera of Proteaceae are hermaphroditic and protandrous, the exceptions being the dioecious southern African genera Aulax and Leucadendron. For hermaphroditic species, the timing of visits by animals to inflorescences is such that they not only acquire pollen from freshly opened flowers but also brush against pollen presenters and stigmas of others that have lost self-pollen and become receptive. Birds and insects (and probably mammals) generally forage in such a way as to facilitate both outcrossing and selfing. Some species are self-compatible, although many require outcrossing if viable seed is to be formed. Regardless of which animals are the major pollen vectors, fruit set is low relative to the number of flowers available, especially in Australian habitats. Functional andromonoecy of the majority of flowers is advanced as the major cause of poor fruit set. The pollination biology and breeding systems of Australian and southern African Proteaceae resemble one another in many ways, partly because of their common ancestry, but also due to convergence. Divergence is less obvious, apart from the dichotomy between dioecious and hermaphroditic genera, and differences in the levels of seed set for Australian and African species. Future studies should concentrate on identifying the most important pollinators for various Proteaceae, the manner in which their visits are integrated with floral development and factors responsible for limiting fruit set.     

Pollinator preference and pollen viability mediated by flower color synergistically determine seed set in an Alpine annual herb

Gentiana leucomelaena manifests dramatic flower color polymorphism, with both blue‐ and white‐flowered individuals (pollinated by flies and bees) both within a population and on an individual plant. Previous studies of this species have shown that pollinator preference and flower temperature change as a function of flower color throughout the flowering season. However, few if any studies have explored the effects of flower color on both pollen viability (mediated by anther temperature) and pollinator preference on reproductive success (seed set) in a population or on individual plants over the course of the entire flowering season. Based on prior observations, we hypothesized that flower color affects both pollen viability (as a function of anther temperature) and pollen deposition (as a function of pollinator preference) to synergistically determine reproductive success during the peak of the flowering season. This hypothesis was tested by field observations and hand pollination experiments in a Tibetan alpine meadow. Generalized linear model and path analyses showed that pollen viability was determined by flower color, flowering season, and anther temperature. Anther temperature correlated positively with pollen viability during the peak of the early flowering season, but negatively affected pollen viability during the peak of the mid‐ to late flowering season. Pollen deposition was determined by flower color, flowering season (early, or mid‐ to late season), and pollen viability. Pollen viability and pollen deposition were affected by flower color that in turn affected seed set across the peak of the flowering season (i.e., when the greatest number of flowers were being pollinated). Hand pollination experiments showed that pollen viability and pollen deposition directly influenced seed set. These data collectively indicate that the preference of pollinators for flower color and pollen viability changed during the flowering season in a manner that optimizes successful reproduction in G. leucomelaena. This study is one of a few that have simultaneously considered the effects of both pollen viability and pollen deposition on reproductive success in the same population and on individual plants.     

Seed set variation in wild Clarkia populations: teasing apart the effects of seasonal resource depletion,pollen quality,and pollen quantity

In habitats where resource availability declines during the growing season, selection may favor early‐flowering individuals. Under such ephemerally favorable conditions, late‐blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation (PL) of seed set and outcrossing rates between early‐ and late‐flowering individuals in two mixed‐mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008–2010, we hand‐pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open‐pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open‐pollinated flowers. Second (2009), we conducted an anther‐removal experiment to estimate self‐pollen deposition. Seed set in Clarkia unguiculata was not pollen‐limited. Clarkia xantiana ssp. xantiana was pollen‐limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator‐dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 – and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules.     

Flowering phenology, display size, and fruit set in an understory dioecious shrub, Aucuba japonica (Cornaceae)

I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.     

Andromonoecy in Solanum lycocarpum A. St. -Hil. (Solanaceae): Floral attributes,visitors and variation in sexual expression over time

Sexual expression in andromonoecious species—those in which a single individual can bear both staminate and hermaphroditic flowers—may vary among reproductive events in the same plant, among individuals and across populations. This variation influences, in turn, the individual contribution of hermaphroditic plants via male and female fitness functions (i.e., Lloydʼs phenotypic gender). However, temporal variation in sexual expression in andromonoecious species and its relationship with seasonal changes in climatic conditions remain poorly understood. Here we analyze floral attributes, visitors and variation in sexual expression in three populations of Solanum lycocarpum A. St. -Hil. Seasonality in the production of floral types, the mating system and floral visitors were also investigated. Hermaphroditic flowers produced more pollen grains, but the pollen of staminate flowers had higher viability. Only hermaphroditic flowers produced fruits, and ovules in staminate flowers were sterile. Solanum lycocarpum is mainly pollinated by large bees with the ability to vibrate flowers. Phenotypic gender varied throughout the year, and the seasonal production of staminate flowers is associated with the local climate. We suggest that the higher and seasonally variable relative abundance of staminate flowers compared to the low and uniform production of hermaphroditic flowers may be explained by (a) the very high energetic costs incurred in producing large fruits, which in turn make hermaphroditic flower production very costly, and (b) the potentially lower energy expenditure of the smaller staminate flowers with reduced pistils and non-viable ovules that allow them to rapidly respond to climate variability.     

FLOWERING,SEX RATIOS,POLLEN-OVULE RATIOS,FRUIT SET,AND REPRODUCTIVE EFFORT OF A DIOECIOUS TREE,MYRISTICA INSIPIDA (MYRISTICACEAE), IN TWO DIFFERENT RAIN FOREST COMMUNITIES

The flowering of Myristica insipida R. Br. was studied in two rain forest communities in northern Queensland. This dioecious, subcanopy tree had a male-biased sex ratio at both study sites. In the lowland population the male-bias could be attributed to males (trees producing staminate flowers) starting to flower at a smaller average size than females (trees producing pistillate flowers). There were no intersexual differences in spacing or distribution within the study sites. Males trees flowered earlier, flowered longer, and produced over twice as many flowers as females during the study season. Although the onset of flowering was rather variable, 18–22 days following heavy rains, most trees had a synchronous period of maximum flowering. Pollination manipulations determined that there was no fruit development without pollination, and that increasing pollen loads resulted in increased fruit set with diminishing effect. Taking into account the sex-ratios and intersexual differences in flower production, the pollen-ovule ratio was calculated to be 16,000–19,000. Male trees were found to expend more energy on flowering than female trees. Open-pollination resulted in 1.0% of female flowers setting fruit. The much greater cost of fruit production resulted in females expending 421% more energy on reproduction than males. Fruit and seed production were judged to be pollination-limited. Nonetheless, this species exhibited several characteristics that are predicted if dioecy evolved by means of sexual selection.     

FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

Reproductive biology in a tropical shrubland of Venezuelan Guayana

Plant sexual systems were evaluated in a tropical shrubland community of Venezuelan Guayana. Of 89 plant species, 73.0% were hermaphrodite, 12.4% monoecious, 10.1 % submonoecious, and 4.5 % dioecious. Temporal variation of sexual expression showed that hermaphrodite species were mostly adichogamous and monoecious, submonoecious species dichogamous (protandrous andprotogynous). Herkogamy was an important mechanism promoting cross-pollination. Of 38 hermaphrodite species, 39.5 % were herkogamous. Plant sexual systems were not associated with life form. Most species were pollinated mostly by large and medium-size bees. Pollinator specificity and pollen load transportation, were higher for monoecious species than for hermaphrodite, submonoecious and dioecious ones. Both pollinator specificity and pollen transportation were not significantly different among representatives of the latter three sexual systems. Plant sexual systems were significantly associated with dispersal syndromes; dioecious species had bird-dispersal syndromes, the other sexual systems mostly anemochory, epizoochory and endozoochory; but many hermaphrodites were anemochorous.     

Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz‐pollinated species with anther dimorphism

Some pollination systems, such as buzz‐pollination, are associated with floral morphologies that require a close physical interaction between floral sexual organs and insect visitors. In these systems, a pollinator's size relative to the flower may be an important feature determining whether the visitor touches both male and female sexual organs and thus transfers pollen between plants efficiently. To date, few studies have addressed whether in fact the “fit” between flower and pollinator influences pollen transfer, particularly among buzz‐pollinated species. Here we use Solanum rostratum, a buzz‐pollinated plant with dimorphic anthers and mirror‐image flowers, to investigate whether the morphological fit between the pollinator's body and floral morphology influences pollen deposition. We hypothesized that when the size of the pollinator matches the separation between the sexual organs in a flower, more pollen should be transferred to the stigma than when the visitor is either too small or too big relative to the flower. To test this hypothesis, we exposed flowers of S. rostratum with varying levels of separation between sexual organs, to bumblebees (Bombus terrestris) of different sizes. We recorded the number of visits received, pollen deposition, and fruit and seed production. We found higher pollen deposition when bees were the same size or bigger than the separation between anther and stigma within a flower. We found a similar, but not statistically significant pattern for fruit set. In contrast, seed set was more likely to occur when the size of the flower exceeded the size of the bee, suggesting that other postpollination processes may be important in translating pollen receipt to seed set. Our results suggest that the fit between flower and pollinator significantly influences pollen deposition in this buzz‐pollinated species. We speculate that in buzz‐pollinated species where floral morphology and pollinators interact closely, variation in the visitor's size may determine whether it acts mainly as a pollinator or as a pollen thief (i.e., removing pollen rewards but contributing little to pollen deposition and fertilization).     

Pollination success in a population of dioecious rain forest trees

Pollination success in female trees was determined for a population of Neolitsea dealbata (R. Br.) Merr., a locally abundant dioecious tree pollinated by small, unspecialized insects in northern Queensland rain forest, Australia. The population consisted of a clustered group of trees with a mean male-to-female distance of 4.5 m and more isolated individuals, including females more than 90 m away from the nearest pollen source. A map of all reproductive trees was produced to determine accurate male-to-female distances. The size of the pollen source available to females was defined as a function of the distance to the nearest ten male trees and their sizes (male neighbourhood index). The rate of pollen movement to females was measured by counting pollen tubes (and the number of tubes per style) in female trees 6 days after the commencement of population flowering. The pollination rate decreased steeply to less than half when the nearest male was only 6.5 m away. Although pollen reached a female 330 m away from the nearest pollen source, only 10% of receptive flowers had been pollinated. The short flowering period (2–3 weeks) combined with the the slow rate of pollen movement means that a large proportion of flowers in isolated trees are unpollinated, confirming an earlier finding that isolated females set fewer fruits than gregarious females. The reliability of pollen transfer to females was determined by quantifying insects and their pollen loads trapped at female trees with a range of male neighbourhood indices. Quantities of insects and pollen were significantly correlated with the size of the male neighbourhood index, indicating a strong density-dependent response by vectors to flowering. Pollen was also collected from insect visitors to non-flowering trees. Females with large male neighbourhood indices received more pollen than non-flowering trees with equivalent male neighbourhood indices. However, when the male neighbourhood indices were small for both female and non-flowering trees, the changces of pollinators encountering female and non-flowering trees were similar, suggesting random movements of pollinators in sparse-flowering sub-populations. The dioecious breeding system, brief, synchronous flowering period, clustered population structure and random, opportunistic foraging behaviour of vectors interacted in a way that reduced reproduction in relatively isolated trees. These results demonstrate a mechanism for differential breeding success between trees in natural populations and emphasize the possible impact of logging regimes on pollen flow between trees. Large interconspecific distances in species-rich environments may have been a factor in the selection for synchronous flowering between trees in outcrossing tree species with generalist insect pollinators.     

Pollen movement to flowering canopies of pistillate individuals of three rain forest tree species in tropical Australia

This study measured the quantities of effective pollen vectors and their pollen loads arriving at the canopies of dioecious tropical rain forest trees in north-east Queensland. Population flowering synchrony, effective pollinator populations and pollen loads transferred between staminate and pistillate trees were compared among three insect-pollinated tree species. All three were visited by a wide range of insects, 75% of which (mostly 3–6 mm long) carried conspecific pollen. Fewer than 8% of individual insects were found to be carrying single-species pollen exclusively and none could be described as specialist pollen foragers. The introduced honeybee carried greater quantities of pollen than any native species but was not necessarily a reliable pollinator. The brief flowering periods in Neolitsea dealbata (3–4 weeks) and Litsea leefeana (4–5 weeks) populations were synchronized among individuals. Flowering in the Diospyros pentamera population extended over 15 weeks and most individuals were in flower for most of this period. Staminate trees began flowering earlier, produced more flowers and attracted relatively more insects than did pistillate trees, suggesting a density-dependent response of pollinators to flowering performance. Pollen was trapped in greater quantities on insects at staminate trees than at pistillate trees. Insect numbers increased at peak flowering periods and Diptera were the most abundant flower visitors. Anthophilous Coleoptera were more numerous at staminate than at pistillate trees in all three tree species populations. Larger quantities of pollen were mobilized during peak flowering times although the greatest quantities were transferred to pistillate canopies towards the end of the population flowering periods. Diptera carried pollen more often to pistillate N. dealbata and L. leefeana trees than did other groups whereas Coleoptera carried pollen more often to pistillate D. pentamera trees. The two contrasting flowering performances in the three tree species are discussed with reference to mechanisms that facilitate pollen transfer between staminate and pistillate trees.