Night and day: Contributions of diurnal and nocturnal visitors to pollen dispersal, paternity diversity, and fruit set in an early-blooming shrub, Daphne jezoensis.

PREMISE: Under uncertain pollinator visit conditions, plants often exhibit long flowering periods and generalized pollination systems. Flowering of the gynodioecious shrub Daphne jezoensis occurs in early spring in cool temperate forests. Pollination by nocturnal moths is expected, given the species' tubular-shaped flowers with sweet fragrance and nectar. However, the effectiveness of nocturnal moths under cool conditions is unknown. We evaluated the relative importance of diurnal and nocturnal visitors as pollinators in early spring. METHODS: We investigated flowering duration, flower visitors, and floral scents in a natural population. We experimentally exposed flowers to visitors only during daytime or nighttime using bagging treatments and evaluated the contributions of diurnal and nocturnal insects to fruit set, pollen dispersal distance, and paternity diversity using 16 microsatellite markers. RESULTS: Female flowers lasted ~3 wk, which was ~8 d longer than the flowering period of hermaphrodites. Various insects, including Coleoptera, Diptera, Hymenoptera, and Lepidoptera, visited the flowers during both daytime and nighttime. Flowers emitted volatiles, such as lilac aldehyde isomers and ß-ocimene, which are known to attract moths. Fruit-set rate in the night-open treatment was similar to or higher than that in the day-open treatment. However, pollen dispersal distance in the night-open treatment was shorter than that in the day-open treatment. Paternity diversity was similar in day-open and night-open treatments. CONCLUSIONS: Early-blooming plants ensure pollen receipt and dispersal by having a long flowering period and using both diurnal and nocturnal flower visitors, suggesting the importance of a generalized pollination system under uncertain pollinator visit conditions.

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.

Restricted female function of hermaphrodites in a gynodioecious shrub, Daphne jezoensis (Thymelaeaceae).

Gynodioecy is the coexistence of hermaphrodites and females in a population. It is supposed to be an intermediate stage in the evolutionary pathway from hermaphroditism to dioecy in angiosperm. Hermaphrodites gain fitness through both seed and pollen production whereas females gain fitness only through seed production. As females spread in a gynodioecious population, sexual selection prompts hermaphrodites to invest in male function and male-biased hermaphrodites prevail. In the gynodioecious shrub Daphne jezoensis (Thymelaeaceae), female frequency is stably around 50% in most populations, and fruit-set rate of hermaphrodites is commonly low. Therefore, D. jezoensis is likely at a later stage in the evolutionary pathway. Female function of hermaphrodites (fruit-set rate, selfing rate, seed size, and germination rate) was assessed in three populations under natural conditions. In order to evaluate the potential seed fertility and inbreeding depression by selfing in hermaphrodites, hand pollination treatments were also performed. Over a 2-year period under natural conditions, 18-29% of hermaphrodites and 69-81% of females set fruit. Across all three populations, the mean fruit-set rate ranged 9.5-49.2% in females and only 3.9-10.2% in hermaphrodites. Even with artificial outcross-pollination, 59-91% of hermaphrodites failed to set any fruit. When self-pollination was performed in hermaphrodites, both of fruit-set and germination rates were decreased, indicating early-acting inbreeding depression. In addition, more than half of the hermaphrodite seeds were produced by selfing under natural pollination, but pollinator service was still required. Totally, hermaphrodites performed poorly as seed producers because of the intrinsically-low fruiting ability and a combination of autogamous selfing and strong inbreeding depression, indicating the absence of reproductive assurance. These results indicate that the mating system of D. jezoensis is functionally close to dioecy.

Size-dependent sex allocation and reproductive investment in a gynodioecious shrub.

In sexually dimorphic plants, resource allocation to reproduction often differs between sex morphs. In gynodioecious species, i.e. coexisting hermaphrodite and female plants within a population, females often produce more fruits than hermaphrodites. Since fruit production is costlier than flower production, hermaphrodites and females may regulate flower and fruit production differently in response to resource availability. To clarify the gender-specific strategies of reproductive allocation, we assessed sexual dimorphism in reproductive traits, size-dependent resource allocation, morphological traits, and photosynthetic capacity in a natural population of a gynodioecious shrub, Daphne jezoensis Hermaphrodites had larger flowers and increased flower number with plant size at a rate greater than females, but showed consistently smaller fruit production. Although females did not increase flower production as much as hermaphrodites did as their size increased, they produced 3.7 times more fruits than did hermaphrodites. Despite a large sexual difference in fruiting ability based on hand-pollination, total resource investment in reproduction (the sum of flower and fruit mass) was similar between sex morphs across plant sizes, and there was little sexual difference in the cost of reproduction, i.e. the negative effect of current reproduction on future reproductive effort, in the natural population. In addition, there were no sexual differences in the resource allocation to vegetative organs (leaf and root mass) and photosynthetic capacity (light response photosynthetic rates). Under natural conditions, pollen limitation strongly restricted the fruit production of females, resulting in similar cost of reproduction between hermaphrodites and females.