The Effects of Inflorescence Size and Flower Position on Biomass and Temporal Sex Allocation in <Emphasis Type="Italic">Lobelia sessiliflora</Emphasis>

To test the prediction of sex allocation theory that plants or flowers high in resource status emphasize the female function, we explored the variation in both biomass (the number of pollen grains and ovules) and temporal (male and female durations) sex allocation among and within plants of protandrous Lobelia sessilifolia in relation to plant size and flower position within plants. Among plants, the mean number of pollen grains and ovules per flower of a plant increased with plant size, whereas the mean P/O ratio (number of pollen grains/number of ovules ratio) decreased with plant size. The mean male duration, the mean female duration, and the mean ratio of male duration/flower longevity per flower of a plant were not correlated with plant size. Thus, large plants emphasized female function in terms of biomass sex allocation, which is consistent with the prediction of size-dependent sex allocation theory. The results for temporal sex allocation, however were inconsistent with the theory. Within plants, the mean number of pollen grains and ovules per flower at each position decreased from lower to upper flowers (early to late blooming flowers) and that of the P/O ratio increased from lower to upper flowers. The mean male duration and the mean female duration per flower decreased from lower to upper flowers, whereas the mean ratio of male duration/flower longevity increased from lower to upper flowers. The population sex ratio changed from male-biased to female-biased. Thus, later blooming flowers emphasized the male function in terms of both biomass and temporal sex allocation, consistent with the sex allocation theory, regarding the change in the population sex ratio.     

The relative importance of architecture and resource competition in allocation to pollen and ovule number within inflorescences of Hosta ventricosa varies with the resource pools

Background and Aims

Allocation of resources to floral traits often declines distally within inflorescences in flowering plants. Architecture and resource competition have been proposed as underlying mechanisms. The aim of the present study is to assess the relative importance of resource competition and architectural effects in pollen and ovule production on racemes of Hosta ventricosa, an apomictic perennial herb.

Methods

Combinations of two defoliation treatments (intact and defoliated) and two fruit-set treatments (no-fruit and fruit) were created, and the roles of architecture and resource competition at each resource level were assessed.

Key Results

Pollen and ovule number per flower increased after defoliation, but pollen to ovule ratio per flower did not change. Pollen, ovules and the pollen to ovule ratio per flower declined distally on racemes at each resource level. In the intact treatment, fruit development of early flowers did not affect either pollen or ovule number of late flowers. In the defoliated treatment, fruit development of early flowers reduced both pollen and ovule numbers of late flowers due to over-compensation caused by defoliation. Late flowers on defoliated fruit racemes produced less pollen than intact fruit racemes but the same number of ovules; therefore, the reduction in pollen number was not caused by over-compensation. In addition, the fruit-set rate of early flowers during flowering was higher in intact racemes than in defoliated racemes.

Conclusions

In flowering plants, the relative importance of architecture and resource competition in allocation to pollen and ovules may vary with the resource pools or the overall resource availability of maternal plants.     

Is increased male flower production a strategy for avoidance of predispersal seed predation in andromonoecious plants?

Floral gender in angiosperms often varies within and among populations. We conducted a field survey to test how predispersal seed predation affects sex allocation in an andromonoecious alpine herb Peucedanum multivittatum. We compared plant size, male and perfect flower production, fruit set, and seed predation rate over three years among nine populations inhabiting diverse snowmelt conditions in alpine meadows. Flowering period of individual populations varied from mid‐July to late August reflecting the snowmelt time. Although perfect flower and fruit productions increased with plant size, size dependency of male flower production was less clear. The number of male flowers was larger in the early‐flowering populations, while the number of perfect flowers increased in the late‐flowering populations. Thus, male‐biased sex allocation was common in the early‐flowering populations. Fruit‐set rates varied among populations and between years, irrespective of flowering period. Fruit‐set success of individual plants increased with perfect flower number, but independent of male flower number. Seed predation by lepidopteran larvae was intense in the early‐flowering populations, whereas predation damage was absent in the late‐flowering populations, reflecting the extent of phenological matching between flowering time of host plants and oviposition period of predator moths. Seed predation rate was independent of male and perfect flower numbers of individual plants. Thus, seed predation is a stochastic event in each population. There was a clear correlation between the proportion of male flowers and the intensity of seed predation among populations. These results suggest that male‐biased sex allocation could be a strategy to reduce seed predation damage but maintain the effort as a pollen donor under intensive seed predation.     

Floral sex allocation and reproductive success within inflorescences of Hosta ventricosa,a pseudogamous apomict

Aims Within inflorescences of sexually reproducing hermaphrodites, the production of ovules, fruits and seeds commonly declines from basal (early-opening) to distal (late-opening) flowers, while pollen production remains constant or only changes slightly, with the result that distal (late-opening) flowers become functionally male. However, few empirical studies have specifically examined whether or not changes in allocation to pollen production actually lead to changes in the number of seeds sired, a more direct measure of male fitness. In pseudogamous apomicts, fitness depends on the number of seeds produced; thus, a contrasting pattern of variation in the pollen-to-ovule (P/O) ratio within inflorescences might be expected.Methods We investigated floral sex allocation and reproductive success within racemes of Hosta ventricosa, a pseudogamous apomictic hermaphrodite possessing flowers that open acropetally. We quantified variations in pollen number, ovule number, the P/O ratio and fruit and seed production, from 2007 to 2011, among flowers within racemes of four populations of H. ventricosa in southwest China. Ecological causes for fruit and seed production were evaluated by observing patterns of pollen deposition, flower removal and supplemental pollination.Important findings Pollen number, ovule number and the P/O ratio declined from basal-to-distal positions in all sampled populations (years). Fruit and seed production decreased distally in most populations (years). Low fruit and seed set of distal flowers was not due to pollen limitation because pollen deposition never declined distally and the low fruit and seed set of distal flowers remained even after supplementary pollination was provided. The flower-removal experiment indicated that inter-fruit competition for resources among flowers was common. The low P/O ratio of distal flowers in H. ventricosa might be favored because they were unable to obtain fitness by donating pollen and siring seeds on other plants. Our study may help to understand the adaptive significance of sex allocation among flowers within inflorescences of sexually reproducing hermaphrodites.     

Effects of generalist herbivory on resistance and resource allocation by the invasive plant,Phytolacca americana

Successful invasions by exotic plants are often attributed to a loss of co‐evolved specialists and a re‐allocation of resources from defense to growth and reproduction. However, invasive plants are rarely completely released from insect herbivory because they are frequently attacked by generalists in their introduced ranges. The novel generalist community may also affect the invasive plant's defensive strategies and resource allocation. Here, we tested this hypothesis using American pokeweed (Phytolacca americana L.), a species that has become invasive in China, which is native to North America. We examined resistance, tolerance, growth and reproduction of plant populations from both China and the USA when plants were exposed to natural generalist herbivores in China. We found that leaf damage was greater for invasive populations than for native populations, indicating that plants from invasive ranges had lower resistance to herbivory than those from native ranges. A regression of the percentage of leaf damage against mass showed that there was no significant difference in tolerance between invasive and native populations, even though the shoot, root, fruit and total mass were larger for invasive populations than for native populations. These results suggest that generalist herbivores are important drivers mediating the defensive strategies and resource allocation of the invasive American pokeweed.     

Synchronized protandry and hermaphroditism in a tropical secondary forest tree, Schefflera heptaphylla (Araliaceae)

Selection favoring avoidance of stigma clogging, pollen discounting, self-fertilization, and other negative effects of self-pollination can produce intricate patterns of intra- and interfloral dichogamy in plants bearing numerous flowers. Here we report an extensive study of the relationships among dichogamy, floral sex allocation (pollen-to-ovule ratios), nectar production, floral visitors, mating system, and fruit set in natural populations of Schefflera heptaphylla, a widespread paleotropical secondary forest tree that produces thousands of flowers in a blooming season. Each tree produces 15?C30 sequentially blooming, paniculate, compound inflorescences. Each compound inflorescence has up to three orders of umbellets, which also bloom sequentially. While hand-pollinations showed that S. heptaphylla was capable of self-fertilization, our observations of thousands of flowers showed that strong intra- and interfloral protandry severely restricts both autogamous and geitonogamous self-pollination. All flowers were bisexual, thus the sexual system of the populations we studied was hermaphroditism. The pollen-to-ovule (P/O) ratios were characteristic of outcrossing species, and P/O ratios of flowers in the last-maturing (third order) umbellets were significantly higher than those in earlier-maturing (first and second order) umbellets. Floral visitors were primarily flies (Chrysomya sp. and Syrphinae sp.) and wasps (Vespula sp. and Eumenes sp.). Flowers produced nectar during both the male (pollen presentation) and female (stigma receptivity) stages of their development, and the volume of nectar production was higher in the female stage. Nevertheless, flowers received fewer visits in the female stage than they did in the male stage, and natural fruit set was low, especially in first and third order umbellets. Fruit set from hand cross- and self-pollinations was significantly higher than natural fruit set, indicating pollen limitation of fruit set. Schefflera heptaphylla has also been reported to be andromonoecious. Both hermaphroditism and andromonoecy are consistent with theoretical predictions for variation in sex allocation among sequentially maturing flowers in protandrous species. Further studies comparing hermaphroditic and andromonoecious populations of S. heptaphylla could elucidate the selective factors affecting sex expression, nectar production, and fruit set in species with numerous flowers displaying both intra- and interfloral dichogamy.     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifolius var. longipes H. T. Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim. var. longipes H. T. Chang, a plant species that only occurs in a few counties in Hunan, Jiangxi and Zhejiang Provinces and with a relatively small number of individuals, is ranked as a second Class endangered species for conservation in China. We have studied the effect of pollen and resources available to female reproduction, and the reproductive mechanism of “excess flowers with low fruit set” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang was discussed. Results are as follows Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. The pollen source rather than pollen numbers significantly affected reproduction of this species. In wild populations, producing one fruit needs about 54.8 flowers, and one satiation seed needs about 6.60 flowers or 83.19 ovules. After fertilizing, which was propitious to flower development, the abortion rate of flower buds was decreasing, but the flowering rate was increasing. The fruit set and seed set was also significantly increasing, while abortion rate of fruit was significantly decreasing. With the increasing percentages of cutting leaves, the fruit set decreased, but the abortion rate of fruit shows no significant differentiation among treatments. After cutting branches that were puny, broken and insectin-fested branches, the flower number seemed to be decreasing, but the fruit set and seed set all increased significantly. After removing some flowers, the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal, whereas fruit set calculated with respect to the initial number of flowers remained constant, and the mean weights of per fruit and per seed all decreased significantly. Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population, while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators. There were very low fruit and seed sets in natural populations of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. Different factors may have interacted to effect a low fruit set. A joint adoption of the “selection abortion hypothesis”, “ovary reserve hypothesis” and “male function hypothesis” seems to be the most likely explanation for the reproductive strategy of “excess flowers with few fruit sets” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang.     

Mate limitation in populations of the endangered Convolvulus lineatus L.: A case for genetic rescue?

In self-incompatible clonal plants, the spread of individual plants can exacerbate mate limitation to the point that it becomes a serious constraint on long-term population persistence, especially in small, isolated populations. In such species, it may be necessary to introduce new genetic material from other populations to restore seed production, a strategy termed “genetic rescue”. In this study we assess the potential pertinence of such genetic rescue in the clonal perennial plant Convolvulus lineatus L., whose populations are often highly reduced in spatial extent and are currently being fragmented by land development projects in Mediterranean France. To do so, we quantify fruit production in a range of populations of different size over four years and perform a series of hand-pollination experiments in natural populations to assess whether fruit set is limited by mate availability. We found that C. lineatus is a self-incompatible species that shows extremely low values of fruit set in natural populations and that a principal cause of this low fruit set is a lack of compatible pollen. This may be primarily due to clonal spread that causes individual populations to be comprised of patches containing one or very few incompatibility types. In small populations fragmented by human activities and which show an absence of fruit production, we thus argue that genetic rescue represents a promising conservation management strategy to avoid inevitable long-term future population decline. We discuss how best to introduce new genetic material into the study populations.     

Environmental Tolerances of Miscanthus sinensis in Invasive and Native Populations

Miscanthus sinensis is a moderately invasive ornamental grass species being considered as a bioenergy species in the USA and elsewhere. In this study, we show the range of environmental conditions tolerated by this species in wild populations in the USA and in Japan. Six naturalized populations in the USA and five native populations in Japan were sampled in summer 2009. In each population, environmental factors (canopy cover and soil fertility) were measured, along with measurements of size and morphology for 30 plants. Relationships between M. sinensis size and environmental variables in the two countries were determined using linear mixed effects models. Results indicated that M. sinensis can tolerate extremely wide variation in soil and climate conditions in the populations we sampled across both ranges, suggesting that it could be successfully grown across a wide distribution in the USA, both intentionally as a bioenergy crop and unintentionally as an escaped invader. Plant size generally responded to different environmental conditions in both ranges, with USA plants being negatively influenced by canopy cover and Japanese plants being positively influenced by soil fertility measures. We recommend caution in growing M. sinensis for bioenergy or ornamental purposes to minimize escape outside of its native range.     

Population dynamics and the ecological stability of obligate pollination mutualisms

Mutualistic interactions almost always produce both costs and benefits for each of the interacting species. It is the difference between gross benefits and costs that determines the net benefit and the per-capita effect on each of the interacting populations. For example, the net benefit of obligate pollinators, such as yucca and senita moths, to plants is determined by the difference between the number of ovules fertilized from moth pollination and the number of ovules eaten by the pollinator's larvae. It is clear that if pollinator populations are large, then, because many eggs are laid, costs to plants are large, whereas, if pollinator populations are small, gross benefits are low due to lack of pollination. Even though the size and dynamics of the pollinator population are likely to be crucial, their importance has been neglected in the investigation of mechanisms, such as selective fruit abortion, that can limit costs and increase net benefits. Here, we suggest that both the population size and dynamics of pollinators are important in determining the net benefits to plants, and that fruit abortion can significantly affect these. We develop a model of mutualism between populations of plants and their pollinating seed-predators to explore the ecological consequences of fruit abortion on pollinator population dynamics and the net effect on plants. We demonstrate that the benefit to a plant population is unimodal as a function of pollinator abundance, relative to the abundance of flowers. Both selective abortion of fruit with eggs and random abortion of fruit, without reference to whether they have eggs or not, can limit pollinator population size. This can increase the net benefits to the plant population by limiting the number of eggs laid, if the pollination rate remains high. However, fruit abortion can possibly destabilize the pollinator population, with negative consequences for the plant population.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

红姜花(姜科)同步大量开花的适应意义

两性花植物同步大量开花增加了个体花展示大小,能吸引更多的传粉者,使植株获得更多的交配机会,但也不可避免地带来不同程度的同株异花授粉,进而对被子植物的性资源分配、花部性状以及雌雄异株的进化等产生重要的影响.本文以姜科红姜花(Hedychium coccineum)为研究对象,通过人工授粉和操控试验、传粉动物的观察、自然结果率和自然居群的大小和密度的调查,探讨红姜花同步大量开花的适应意义.结果表明:(1)红姜花的穗状花序由57.33±1.68(n=30)个蝎尾状聚伞小花序组成,每个小花序具3.8±0.15(n=30)朵花,同花序上所有的小花序同步开花,不同小花序内的花按轮次同步开放,使花序在整个开花期间保持有数十朵花同时开放,花序水平上的花展示极为显著;(2)红姜花为自交亲和植物,但不存在自动自花授粉,自然居群存在严重的传粉者限制,其结实也存在着资源限制;(3)多型蓝凤蝶(Papilio memnon)、小矩翅粉蝶(Dercas lycorias)和黑角尖翅粉蝶云南亚种(Appias indra aristoxemus)是红姜花的有效传粉者,3种蝴蝶对红姜花不同大小花序的访问频率都显著不同,随着花序花展示大小的增加其访问频率也显著增加;(4)红姜花同一个体小花序内不同轮次花之间的胚珠数随轮次显著减少,而花粉数前3轮随轮次显著增加.胚珠数的递减及结实存在着资源限制,说明红姜花一个花序开大量的花并不是为了增加结实,而花粉数前3轮的递增,有利于花粉的分散输出,提高个体的雄性适合度.     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

SEX RATIO,COMPONENTS OF REPRODUCTION,AND POLLEN DEPOSITION IN LINDERA BENZOIN (LAURACEAE)

Sex ratio; flower, pollen, ovule, and fruit production; pollen deposition; and pollinator abundance were measured in populations of the dioecious shrub Lindera benzoin occurring in sun and shade habitats. Sex ratio was 1:1 in both the sun and shade. Flower production was greater in the sun than in the shade, and greater for male plants than for females in both habitat types. The population level ratio of pollen to ovules (P/O) was greater in the shade than in the sun, but pollinator abundance was a more important determinant of pollen deposition than P/O. Although 30%–70% of stigmas received no pollen, an even larger proportion of flowers did not set fruit, and fruit set was greater in the sun than in the shade. Increased illumination due to periodic gap formation may play an important role in the reproductive ecology of this understory shrub by affecting patterns of allocation toward reproduction in male and female plants.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

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.     

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.