Masting and insect pollination in the dioecious alpine herb aciphylla

Abstract

Aciphylla species (wild spaniard/speargrass) are an iconic component of the Australasian high country flora, but their reproductive system is enigmatic. They are insect-pollinated dioecious mast seeders (synchronous highly variable seed production), which seems maladaptive. The resource supply to pollinators is highly variable, yet dioecious plants are dependent on pollinators, and dioecious masting requires male and female plants to flower synchronously. Floral display in Aciphylla is relatively large, with tall inflorescences bearing thousands of flowers, suggesting that plants would not have the resources to produce such large stalks every year. But why do they have such huge inflorescences in the first place? I tested whether pollinator attraction is providing an economy of scale which favours intermittent production of very large inflorescences, by manipulating floral display size during a high-flowering year and measuring insect visitation rates and seed set (female reproductive success). Using space-for-time substitution and selective removal of male inflorescences, I also tested whether female seed set was affected by distance to flowering male plants (i.e. changes in local pollen availability) to see if flowering asynchrony would reduce pollination success. Bags were used to exclude pollination by insects and test for wind pollination, and hand pollination was done to test for pollen limitation. Insect surveys suggest that Aciphylla has a generalist pollination system (to avoid satiating a specialist pollinator during 'mast' years'). Male inflorescences received significantly more visits than females, and some seeds were set inside bags (although only 20-30%), suggesting wind pollination may occur at low levels. Seed set rate was higher for taller inflorescences with greater flowering length in A. aurea but tall inflorescences with excess flowers led to a decrease in seed set rates in A. scott-thomsonii. Hand pollination significantly increased seed set rates although these effects were not as large as expected (e.g. 10% increases from natural to hand-pollinated inflorescences were typical). There was no evidence for resource limitation in any species. Female plants in dense flowering populations had higher seed set rates, and individual floral display size in females was particularly important when females were 'isolated' from males. Insect visitation rates were generally higher on inflorescences with a larger floral display, suggesting that display size is important for pollinator attraction. Overall, these results suggest that the pollinator-attraction benefits of such a large floral display (at both the plant and population level) are possibly providing an economy of scale, although the relative effects are small.