Premise Floral stalk height is known to affect seed dispersal of wind‐dispersed grassland species, but it may also affect the attractiveness of flowers and fruits of animal‐pollinated and animal‐dispersed plants. Stalk height may thus be responsive to selection via interactions with both mutualist pollinators and seed dispersers, but also antagonist florivores and seed predators. In this study, we aimed to determine the effect of pollinators and seed predators on selection on floral stalk height in the insect‐pollinated and wind‐dispersed, alpine, andromonoecious herb Pulsatilla alpina, whose flowers also vary in their sex allocation and thus in the resources available to both mutualists and antagonists. Methods We measured the resource status of individuals in terms of their size and the height of the vegetation surrounding plants of P. alpina at 11 sites. In one population, we recorded floral stalk height over an entire growing season and investigated its association with floral morphology and floral sex allocation (pistil and stamen number) and used leaf‐removal manipulations to assess the effect of herbivory on floral stalk height. Finally, in four populations, we quantified phenotypic selection on floral stalk height in four female components of reproductive success before seed dispersal. Results Stalk height was positively associated with female allocation of the respective flower, the resource status of the individual, and the height of the surrounding vegetation, and negatively affected by leaf removal. Our results point to disruptive selection on stalk height in terms of both selection differentials and selection gradients for fertilization, seed predation, and seed maturation rates and to positive selection on stalk height in terms of a selection differential for mature seed number. Conclusions Stalk height of P. alpina is a costly trait that affects female reproductive success via interactions with both mutualists and antagonists. We discuss the interplay between the resource status and selection imposed on female reproductive success and its likely role in the evolution of sex‐allocation strategies, especially andromonoecy.
The optimal life history and sex allocation of perennial hermaphrodites should depend on both their size and the relative costs and benefits of reproducing through male versus female functions. Theory predicts that insect-pollinated perennials should increase their allocation to female function with size, while the 'mating environment' hypothesis predicts that allocation to male function should track mating opportunities over the course of flowering. We test these two predictions by inferring male and female reproductive success in the protogynous perennial herb Pulsatilla alpina by tracking the patterns and dynamics of sex allocation over time for marked individuals over a range of sizes. We found that small individuals tend to produce only male flowers and that both small and larger individuals produced male flowers at the beginning of the flowering season when mating opportunities were high. By considering within-population variation in life history and phenology jointly rather than separately, and by considering both the tradeoff costs and benefits of allocation to male versus female functions, our results provide new insights into the evolution of both gender diphasy and andromonoecy in perennial plants that are constrained by a dichogamous flowering strategy.
Flowering phenology of alpine plant communities and seasonal dynamics of flower visitors have been scarcely studied in the tropical/subtropical alpine regions. We report flowering phenology, flower production, and flower-visiting insects in the alpine site of central Taiwan. Throughout the research period (2017–2018), we recorded flowering phenology of 130 plant species, flower production of 81 species, and 15,127 insects visiting alpine flowers. Most of the alpine plants were visited by dipteran insects and/or hymenopteran insects. The seasonal patterns of flowering were more apparent in bee-visited plants compared to fly-visited plants in which the flowering of bee-visited plants clearly increased as the season progressed. About 63% of flower visitors were dipteran insects (syrphid and non-syrphid flies), and 30% were hymenopteran insects (mostly bumble-bee workers). Although the seasonal trend in fly abundance was less clear between years, bumble-bee abundance consistently increased in the middle to late seasons, reflecting colony development. There was a positive correlation between bee abundance and the number of flowering species of bee-visited plants, but there was no correlation between fly abundance and the number of flowering species of fly-visited plants throughout the season. These results suggest that the flowering phenology of subtropical alpine communities is influenced by the seasonal availability of pollinators. Bumble bees, syrphid flies, and non-syrphid flies had wide ranges of foraging flowers, but their niche overlap was relatively small. Because cold-adapted bumble bees are threatened by climate change in Taiwan, plant–pollinator interactions may be disturbed by global warming.
Aeschynanthus (Gesneriaceae), a genus comprising approximately 160 species in subtropical Southeast Asia, has red, tubular flowers, typical of a sunbird pollination syndrome. A. acuminatus, the species that is distributed extending to the northern edge of the genus, where the specialized nectarivorous sunbirds are absent, possesses reddish-green flowers and a wide-open corolla tube, flowering time shifts from summer to winter and the species achieves high fruiting success. This atypical flower led us to investigate the pollination biology of this species. Three species of generalist passerines, Grey-cheeked Fulvetta (Alcippe morrisonia, Sylviidae), White-eared Sibia (Heterophasia auricularis, Leiothrichidae) and Taiwan Yuhina (Yuhina brunneiceps, Zosteropidae), were recorded visiting A. acuminatus flowers. Pollination effectiveness was quantified via conspecific pollen presence on stigmas and natural fruit set. The significantly high natural fruit set (60%) and conspecific pollen transfer rate (94%) indicate high reproductive success facilitated by the accurate pollen placement on the birds. The existence of copious (61 µL) and highly diluted (7%) hexose-dominant nectar, together with a major reflectance peak of corolla lobe in the long-wavelength red color spectrum, is consistent with the pollination syndrome of generalist passerines. The high pollination effectiveness of A. acuminatus due to the recruitment of generalist passerines as pollinators, and the specializations of floral traits to match generalist bird pollination, appear crucial in the successful colonization on islands such as Taiwan that lack specialized bird pollinators.
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