Pollination biology and seed production of dioeciousCaryodendron orinocense (Euphorbiaceae) in a plantation in coastal Ecuador

Caryodendron orinocense Karst. (Euphorbiaceae) is an important neotropical, dioecious crop tree providing oil—and protein-rich seeds. Increases in size and number of leaves, flowers, and fruits were estimated for 100 trees in a plantation in costal Ecuador. Height increase was one meter per year—female trees grew less than non-flowering and male individuals. Trees matured at about 4 m height, but flowering and production of fruits remained low until tree height exceeded 8 m. Flowering and leaf production started at the beginning of the rainy season, and flowers were pollinated by large flies. Foliage damage did not seriously reduce leaf number nor fruit production. The plantation studied yielded 150 kg seeds per hectare per year. A plantation with 10 m high trees and a sex ratio of nine females per male will produce 500 kg seeds per hectare per year after about 12 years. Production will increase much more in older plantations.     

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.     

Some pollinators are more equal than others: Factors influencing pollen loads and seed set capacity of two actively and passively pollinating fig wasps

The nursery pollination system of fig trees (Ficus) results in the plants providing resources for pollinator fig wasp larvae as part of their male reproductive investment, with selection determining relative investment into pollinating wasps and the pollen they carry. The small size of Ficus pollen suggests that the quantities of pollen transported by individual wasps often limits male reproductive success. We assessed variation in fig wasp pollen loads and its influence on seed production in actively pollinated (Ficus montana) and passively pollinated (Ficus carica) dioecious fig trees.The ratios of number of male flowers on number of female flowers in a glasshouse-maintained F. montana population were highly variable. When fig wasps were introduced into receptive female figs, the resulting seed numbers were strongly linked to the numbers of pollinators that had been seeking access to pollen, relative to the number of anthers in their natal figs. In F. carica estimates of the amounts of pollen produced per fig and the quantities of pollen carried by emerging fig wasps suggest that less than 10% of the pollen is transported. Pollinators of F. carica that emerged earlier from figs carried more pollen, and also generated more seeds when introduced into receptive female figs.We show here that all pollinators are not equally valuable and producing more pollinators is not necessarily a good option in terms of Ficus male fitness. Previous results on F. montana figs showed that only around half of the flowers where pollinators lay eggs produced adult offspring. The amount of pollen collected by young female fig wasps may be a major determinant of their reproductive success.     

Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea (Lamiaceae)

Summary Pollen movement is often restricted in natural populations, and insufficient pollination is a potential constraint on sexual reproduction in outcrossing species. Seed-set should decrease with increased distance from the pollen source in outcrossing plants. This prediction was tested using females of the clonal, gynodioecious herb Glechoma hederacea in three natural populations. In controlled pollinations, both hermaphrodites and females had similar high percentages of fruit-set and seed-set. In a natural population where a female clone was isolated from the nearest hermaphroditic clone by c. 100 m, fruit-set was low (1%). In another population where hemaphroditic clones were rare and female clones had a patchy distribution, fruit-and seed-set in females were pollen-limited and decreased with increased distance from the nearest pollen source. The estimated mean pollen dispersal distance was 5.9 m when calculated on fruit-set and 5.3 m when calculated on seed-set. The most frequent pollinators were bumblebees. The mean and median distances moved by pollinators between ramets were 0.13 m and 0.05 m. In a third population where female clones were isolated from the nearest hermaphrodites by more than 200 m, fruit-set was 0%. After introduction of 16 hermaphroditic ramets in the center of the female clone, fruit-set varied between 0% and 100% in individual female ramets. Fruit-set decreased with increased distance from the pollen source. The mean and median pollen movement distances were 1.06 m and 0.54 m.     

Limitations to Reproductive Success in the Dioecious Tree Rhamnus davurica

The reproductive success of a female plant in a dioecious species may be affected by pollen limitation and resource limitation. This study presents evidence that the reproductive success of the dioecious understorey tree species, Rhamnus davurica, is affected by the distance to the nearest male. The sex ratios were female-biased, although showing fluctuations in the three years of conducting the study. The mortality rate of females was higher than that of males indicating a trade-off between reproduction and survival. Altogether 49 females, designated as “focal females”, were randomly selected for monitoring their reproductive status between April and October in 2010. But successful reproduction (meaning that the flowering female trees had fruit in the fruiting season) was observed only in 28 females in 2011 and 16 females in 2012. The method of path analysis was applied to determine the effect of topography, local competition and proximity to the nearest male on the fruit set of the females. In the three years of the study, elevation, competition and female size had no significant effect on the fruit set. The distance to the nearest male, however, had a significant effect on fruit set. Number of fruits and fruit set were decreased with increasing distance to the nearest male. It was possible to estimate maximum fruit set, based on the comparatively large dataset. The number of fruits and the fruit set are exponentially related to the distance to the nearest male and the relationships are described by an exponential model. The results of this study support the importance of pollen limitation on the reproductive success in Rhamnus davurica.     

Intersexual mimicry and imperfect deceit of a threatened aquatic herb Ottelia acuminata

Mimicry of non-rewarding flowers to rewarding flowers has been accepted as a strategy to improve pollination success in angiosperms. It has been proposed that this mechanism depends on whether potential pollinators can discriminate between the flowers. In this study, the intersexual mimicry and deceit pollination were studied in a threatened dioecious aquatic herb, Ottelia acuminata. Its female flowers resemble male flowers in morphology and odor compounds, to avoid discrimination by pollinators and outcompete male flowers in attracting the pollinators using stronger scents and bigger flowers. However, an obvious visit bias of its pollinator (Apis cerana) to male flowers was detected, suggesting that bees can distinguish the rewarding males from non-rewarding females. Although the deceit was not successful, pollination was not seriously undermined because pollen limitation was found to be low in the sampled natural population. We speculate that, due to “accidental” visits on female flowers and “mistake” pollinations, pollen limitation could be mitigated by a high frequency of pollen donors, and is correlated with the size and sex ratio of a population. Ottelia acuminata is a threatened dioecious aquatic herb. We suggest that developing multi-stakeholder coalitions should be encouraged to save the threatened edible and ornamental plant species in China. We hope this study could provide new insights into understanding of the role of intersexual mimicry in other flowering plants.     

Impact of plant flowering phenology on the cost/benefit balance in a nursery pollination mutualism,with honest males and cheating females

This study documents the flowering phenology and its potential consequences on a nursery pollination mutualism between a dioecious plant, in which honest male plants, but not cheating females, allow the specific pollinator to reproduce within inflorescences. Very few pollinators were found to emerge during plant anthesis, leading to a low (if any) potential benefit through pollen dispersal. This opens the question why male plants do not also cheat their pollinators. Female plants flowered late in the season, when many males had just achieved their own anthesis, which increased the efficiency of pollen transfer. Finally, some late‐flowering males reached their anthesis simultaneously with females, which open the possibility for pollinator to choose between honest males and cheating females. Nevertheless, female plants were found to produce fruits, even though fruit production was limited by pollen (and pollinator) supply, meaning that cheating was not entirely retaliated by the mutualistic partner.     

Impact of balsam fir flowering on pollen and foliage biochemistry in relation to spruce budworm growth, development and food utilization

The impact of balsam fir (Abies balsamea (L.) Miller) flowering on nutritional and allelochemical quality of pollen, current-year and one-year-old foliage is studied in relation to spruce budworm (Choristoneura fumiferana Clem.) (Lepidoptera: Tortricidae) growth, development and utilization of food and nitrogen. In the laboratory, using fresh food from the field, we simulated conditions of low larval population density, in which there is no current-year foliage depletion during the spruce budworm feeding period. Similarly, we simulated conditions of high larval population density when current-year foliage depletion occurs.Because of the high nutritive value of pollen (high amounts of amino acids and minerals, especially nitrogen; low monoterpene content), insects from flowering trees reached the fifth instar five days earlier than those from non-flowering trees, and had heavier dry- and nitrogen-weights at the beginning of the fifth instar. At budbreak, switching from pollen to current-year foliage negatively affected conversion efficiencies and digestibilities of food and nitrogen (AD; ADN; ECDN; ECI; ECIN). The switch from pollen to new foliage had a detrimental impact on fifth-instar survival and on newly-moulted sixth-instar dry- and nitrogen-weights. Moreover, during the fifth instar, balsam fir flowering reduced the nutritive value of current-year foliage, which in turn, might have contributed to the reduced larval growth. Nevertheless, during the sixth instar, balsam fir flowering affected the biochemistry of current-year foliage in ways that enabled larvae to compensate for their low fifth-instar biological performance; larvae also managed to reach pupal dry weight similar to larvae reared on non-flowering trees. Current-year foliage from flowering trees contained less nitrogen, total soluble sugars and total monoterpenes. Those foliar characteristics enabled larvae to increase food and nitrogen consumption rates (RCR; RNCR), because of lower repellency and/or post-ingestional feedback from monoterpenes.As for current-year foliage, balsam fir flowering reduced nitrogen, total soluble sugar and total monoterpene contents in one-year-old foliage during the sixth-instar feeding period. These characteristics enabled sixth-instar larvae, fed on old foliage from flowering trees, to have high relative food and nitrogen consumption rates (RCR; RNCR). Larvae were also able to reach higher relative growth rates (RGR) and relative nitrogen accumulation rates (RNAR) compared to larvae reared on one-year-old foliage from non-flowering trees. Finally, larvae on flowering trees had pupal dry weight similar to those from non-flowering trees, but reached the adult stage nine days earlier.Regardless the foliage type consumed by spruce budworm larvae during the sixth instar, pollen consumption during early larval stages reduced total development time, and thus exposure time to natural enemies. This phenomenon might increase larval survival. Balsam fir flowering changed the biochemistry of one-year-old and current-year foliages, but did not affect pupal dry weights of larvae reared on flowering trees compared to those reared on non-flowering trees. Thus, at low population density, spruce budworm populations in balsam fir flowering stands might be favoured over those in balsam fir non-flowering stands. In addition, when larvae consumed one-year-old foliage during the entire sixth instar, those on flowering trees are probably favoured over those on non-flowering trees. However, because flowering trees produce less new foliage than non-flowering trees, current-year foliage depletion may occur earlier on flowering trees than on non-flowering trees. Thus, at similar larval population density, larvae on flowering trees might have to feed on one-year-old foliage earlier than those on non-flowering trees. In that case, spruce budworm populations on non-flowering stands would be favoured over those on flowering stands.     

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.     

Pollination of Schisandra henryi (Schisandraceae) by female, pollen-eating Megommata species (Cecidomyiidae, Diptera) in south-central China

BACKGROUND AND AIMS: The mutualistic interaction between insects and flowers is considered to be a major factor in the early evolution of flowering plants. The Schisandraceae were, until now, the only family in the ANITA group lacking information on pollination biology in natural ecosystems. Thus, the objective of this research was to document the pollination biology and breeding system of Schisandra henryi. METHODS: Field observations were conducted in three populations of S. henryi and the floral phenology, floral characters and insect activities were recorded. Floral fragrances were sampled in the field and analysed using TCT-GC-MS. Floral thermogenesis was measured with a TR-71U Thermo Recorder. Pollen loads and location of pollen grains on insect bodies (including the gut) were checked with a scanning electron microscope and under a light microscope. KEY RESULTS: Schisandra henryi is strictly dioecious. Male flowers are similar to female flowers in colour, shape, and size, but more abundant than female flowers. The distance between tepals and the androecium or gynoecium is narrow. Neither male nor female flowers are fragrant or thermogenic. Schisandra henryi is pollinated only by adult female Megommata sp. (Cecidomyiidae, Diptera) that eat the pollen grains as extra nutrition for ovary maturation and ovipositing. Both male and female flowers attract the pollinators using similar visual cues and thus the female flowers use deceit as they offer no food. CONCLUSIONS: Schisandra henryi exhibits a specialized pollination system, which differs from the generalized pollination system documented in other ANITA members. Pollen is the sole food resource for Megommata sp. and the female flowers of S. henryi attract pollinators by deceit. This is the first report of predacious gall midges utilizing pollen grains as a food source. The lack of floral thermogenesis and floral odours further enforces the visual cues by reducing attractants for other potential pollinators.     

Evidence for intersexual chemical mimicry in a dioecious plant

The dioecious Mediterranean fig, Ficus carica, displays a unique phenology in which males sometimes bloom synchronously with females (in summer), and sometimes not (in spring). Ficus carica is engaged in an obligatory mutualism with a specific pollinating wasp, which reproduces only within figs, localising them by their specific scents. We show that scents emitted by male figs show seasonal variation within individual trees. Scents of summer male figs resemble those of the co-flowering females, and are different from those of the same male trees in spring, when female figs are absent. These differences hold even if only compounds electrophysiologically active for pollinators are considered. The similar scents of summer males and females may explain why the rewardless females are still pollinated. These results offer a tractable model for future studies of intersexual chemical mimicry in mutualistic pollination interactions.     

Pollination in a cryptically dioecious plant Decaspermum parviflorum (Lam.) A. J. Scott (Myrtaceae) by pollen-collecting bees in Sulawesi, Indonesia

Decaspermum parviflorum is a common shrub or treelet in clearings in eastern Sulawesi, Indonesia. It is widely distributed in SE Asia. It produces numerous white flowers and small blue-black fruits. Individual shrubs were of two types in 1:1 ratio, one bearing staminate flowers, the other bearing flowers with both stamens and carpels. The 'hermaphrodite' flowers were shown to have sterile pollen so the species is functionally dioecious. The flowers and floral sprays of the two sexes differed in several measurements: the male flowers had more anthers and were larger and more numerous, making the corporate visual image of male sprays more than twice as large as that for sprays of female flowers. Flowers, individuals of which last only one day, opened only every second day, when all bushes flower synchronously. When the anthers burst in the morning, pollen and sterile pollen was collected vigorously by a variety of bees, mostly Apis dorsata and Nomia spp. Most of the pollen was collected within 1 h of anthesis. Anthesis in males takes place about 20 min earlier than in females; insect visitation follows this pattern. The minute quantity of nectar was collected by only few insects and mostly after the pollen had gone, if at all. Fruits were eaten by birds. The species displays many features typical of dioecious tropical plants with the unusual feature of pollen being the main food reward for pollinators. It was pollinated effectively in Sulawesi.     

Pollinators entering female dioecious figs: why commit suicide?

In the dioecious fig/pollinator mutualism, the female wasps that pollinate figs on female trees die without reproducing, whereas wasps that pollinate figs on male trees produce offspring. Selection should strongly favour wasps that avoid female figs and enter only male figs. Consequently, fig trees would not be pollinated and fig seed production would ultimately cease, leading to extinction of both wasp and fig. We experimentally presented pollinators in the wild (southern India) with a choice between male and female figs of a dioecious fig species, Ficus hispida L. Our results show that wasps do not systematically discriminate between sexes of F. hispida. We propose four hypotheses to explain why wasp choice has not evolved, and how a mutualism is thus maintained in which all wasps that pollinate female figs have zero fitness.     

Reproductive biology and pollination ecology of Triplaris gardneriana (Polygonaceae): a case of ambophily in the Brazilian Chaco

• Triplaris gardneriana (Polygonaceae) is a dioecious pioneer tree reported as insect‐pollinated, despite possessing traits related to anemophily. Here, we analyse the possible roles of insects and wind on the pollination of this species to establish whether the species is ambophilous.
• We carried out observations of floral biology, as well as on the frequency and behaviour of pollinators visiting flowers in a population of T. gardneriana in the Chaco vegetation of Brazil. We conducted experimental pollinations to determine the maternal fertility of female plants and whether they were pollen‐limited, and we also conducted aerobiological experiments to provide evidence of how environmental factors influence atmospheric pollen dispersal.
• The population comprised an area of approximately 152.000 m² and was composed of 603 female and 426 male plants (sex ratio = 0.59:0.41). We observed 48 species of insects visiting flowers of T. gardneriana, of which the bees Scaptotrigona depilis and Apis mellifera scutellata were the most effective pollinators. We recorded pollen grains dispersed by wind on 74% of the glass slides placed on females, located at different distances (1–10 m) from male plants.
• Airborne pollen concentration was negatively correlated with relative humidity and positively correlated with temperature. Our observations and experimental results provide the first evidence that T. gardneriana is an ambophilous species, with pollen dispersal resulting from both animal and wind pollination. This mixed pollination strategy may be adaptive in T. gardneriana providing reproductive assurance during colonisation of sites with different biotic and abiotic conditions.

     

Size distribution,growth and inter-year variation in sex expression of Bischofia javanica,an invasive tree

Flowering activity and sex expression of Bischofia javanica Blume were investigated for 3 years. B. javanica is an invasive dioecious tree of subtropical forests on the Bonin Islands in the western Pacific of Japan. The sex ratio showed a significant male bias (1.25-2.33). Smaller trees were significantly male biased, whereas larger trees showed no significant difference in sex expression, suggesting that males tend to be more precocious in sexual reproduction. We found evidence for sex changes in B. javanica; these have not been reported previously. Most of the 1,653 census trees remained non-flowering (58.1 %); 3.7 % of them showed sex changes, and the percentage of trees repeatedly flowering as males and females was 10.5 and 3.4 %, respectively. Sex changes were observed in both directions but a larger percentage of male trees became female. Flowering frequency and sex expression were significantly related to tree size (i.e. diameter at breast height). Over the 3 years, trees that were consistent females were the largest; inconsistent trees (switching sex between years) were intermediate in size, whereas consistent males were the smallest. There were no significant differences in relative growth rate (RGR) among trees of different sex or flowering frequencies. These results suggest that the maintenance of female reproduction is not related to changes in RGR of diameter but to flowering frequency or the reversal to the male form, dependent upon the internal resource status of individual trees.     

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.     

Pollen limitation and fruit abortion in a declining rare tree,the Eurasian yew (Taxus baccata L.): A reproductive cost of ecological marginality

Populations of rare tree species such as the dioecious and anemophilous yew (Taxus baccata) are especially prone to extinction if they occupy marginal habitats. This is the case of yew populations growing in Mediterranean mountains under dry climate and a severe anthropogenic disturbance regime, which show insufficient regeneration. We examined two of the likely mechanisms driving this population decline. The study was conducted in a central Spanish yew population in 2005 and 2006. On one hand, we tested for the effect of the amount of pollen received by female flowers by means of a pollen addition experiment. On the other hand, we tested for the effect of initial fruit set on abortion and mature fruit production. Finally, we evaluated the effect of the spatial arrangement of male and female yew trees on realized fecundity. The success of fertilization and fruiting in the experimental population was pollen-limited. Female trees aborted ca. 70% of the fruit set in both years and treatments. Fruit set was significantly influenced by the females' neighbourhood, with fecundity decreasing exponentially with distance to the nearest male in both years. Overall, our results indicate that factors associated with ecological marginality have a strong influence on yew performance.     

Relative importance of diurnal and nocturnal pollinators for reproduction in the early spring flowering shrub Stachyurus praecox (Stachyuraceae)

Generalized pollination systems may be favored in early spring flowering plants, as during this period pollinator activity is unpredictable. Many previous studies have concentrated on the importance of diurnal visitors in pollination, and consequently, information on the contribution of nocturnal visitors to pollination in early spring is limited. This study was conducted to evaluate the relative importance of diurnal and nocturnal pollinators in the early spring flowering dioecious shrub Stachyurus praecox (Stachyuraceae), in two temperate forests in central Japan. Visitors to the female and male flowers were observed during day and night, and their relative contributions to seed set were compared. The pollinator observations revealed that the diurnal and nocturnal insects visited both male and female flowers, and that the main flower visitors were diurnal small bees and flies as well as nocturnal settling moths. The diurnal and nocturnal flower visitors also acted as pollinators, as the pollen grains of S. praecox were attached to the insects collected from the female flowers. Pollination experiments demonstrated that the contributions of diurnal pollinators to the seed set were higher than those of the nocturnal pollinators. The results of this study indicate that S. praecox has a generalized pollination system, comprising both diurnal insects and nocturnal settling moths. Although the roles of diurnal insects are more important in the pollination of S. praecox, nocturnal settling moths may have a complementary role in early spring.     

When looks can kill: the evolution of sexually dimorphic floral display and the extinction of dioecious plants

Dioecious plants (with separate male and female individuals) more often have drab, inconspicuous flowers than related bisexual plants. Models indicate, however, that similar conditions favour the evolution of showy floral displays in dioecious and bisexual plants. One difference, however, is that dioecious plants may evolve floral displays that are sexually dimorphic. We show that males are more likely to evolve showy flowers than females in animal-pollinated plants, especially when pollinators are abundant. We demonstrate that this dimorphism places showy dioecious plants at a much higher risk of extinction during years of low pollinator abundance because pollinators may fail to visit female flowers. The higher extinction risk of showy dioecious plants provides an explanation for the fact that dioecious plants that do persist tend to have inconspicuous flowers and are more often wind pollinated. It may also help explain why dioecious plants are less species-rich than related bisexual plants.     

Complex patterns of mating revealed in a Eucalyptus regnans seed orchard using allozyme markers and the neighbourhood model

The neighbourhood model apportions offspring of individual mother plants to self-fertilization, outcrossing to males within a circumscribed area around the mother plant (the neighbourhood), and outcrossing to males outside the neighbourhood. Formerly the model was applied only to haploid pollen gametes in the offspring of conifers, but is extended so that it can be used with genotypic data from diploid offspring of both angiosperms and gymnosperms. In addition, it is shown that the mating parameters can be estimated without independent estimates of allele frequencies in the pollen pools outside the neighbourhood; thus the model might be applied effectively to natural populations exposed to unknown external pollen sources. Parameters of the neighbourhood mating model were estimated for a 10-year-old seed orchard population of the insect-pollinated tree, Eucalyptus regnans, in southeast Australia, which contained a mixture of two geographical provenances (Victoria and Tasmania). The mating patterns revealed were complex. Crosses between trees of the same provenance occurred three times more often than crosses between trees of different provenances. Levels of self-fertilization and patterns of mating within neighbourhoods were influenced by provenance origin, crop fecundity and orchard position (central vs. edge) of mother trees. Gene dispersal, however, was extensive, with approximately 50% of effective pollen gametes coming from males more than 40 m away from mother trees (average distance between neighbouring trees was 7.4 m). Thus, insect pollinators are efficient promoters of cross-fertilization in this orchard, with the result that the effective number of males mating with each female is large.