Horizontal orientation facilitates pollen transfer and rain damage avoidance in actinomorphic flowers of Platycodon grandiflorus.

In zoophilous plants, floral orientation evolved under both biotic and abiotic pressure to enhance pollination success. However, the adaptive significance of horizontal orientation in radially symmetrical (actinomorphic) flowers remains largely unknown, although that of bilaterally symmetrical flowers has been well studied. We experimentally altered floral angle in a population of insect-pollinated Platycodon grandiflorus flowers to examine the effects of floral orientation on pollinator behaviour, pollination success and pollen rain damage avoidance. To further investigate the potential pollen damage by rain, we obtained past precipitation records for the study area during the flowering season, and experimentally tested P. grandiflorus pollen damage by water. Horizontally oriented flowers received more pollinator visits and had pollen grains on the stigma in male and/or female phases than downward- and/or upward-oriented flowers and avoided pollen damage by rainfall better than upward-oriented flowers. A pollen germination experiment showed that approximately 30% of pollen grains burst in distilled water, indicating that pollen damage by rainfall may be significant in P. grandiflorus. Our field experiments revealed that upward-oriented flowers cannot avoid pollen damage by rainfall during the flowering period, and that both upward- and downward-oriented flowers experience pollinator limitation in female success. Therefore, horizontal flower orientation appears to be adaptive in this insect-pollinated actinomorphic species that blooms during the rainy season.

Covering and shading by neighbouring plants diminish pollinator visits to and reproductive success of a forest edge specialist dwarf species.

The pollination and reproductive success of flowering plants can be negatively influenced in various ways by neighbouring heterospecific plants, such as resource competition and reproductive interference. We hypothesized that covering together with shading by neighbouring plants may reduce pollinator visits to and reproductive success of plants by reducing floral attractiveness and pollinator activity and by interrupting flower access, respectively. To test this hypothesis, we examined whether shaded and covered flowers suffered from pollinator limitation and low reproductive success in a population of the dwarf herb Lithospermum zollingeri, which co-exists with woody and herbaceous plants in anthropogenically maintained forest edge meadows. Here, shaded and covered flowers were defined as those beneath the shade of the woods and those whose front portion was covered by any vegetative part of neighbouring plants, respectively. The shaded and covered flowers were visited by significantly fewer pollinators than sunlit and open flowers in the field. However, three major pollinator species responded differently to shading and covering. Significant pollen limitation reduced seed set in covered flowers, and shaded flowers produced fewer seeds. Pollen removal from the anthers was not influenced by shading or covering. Our study demonstrates the negative effects of covering on pollinator visits and seed production. It also elucidates the negative effects of shading on reproductive success in L. zollingeri, which depends on managed semi-natural conditions. Land management abandonment, which has increased shaded and covered conditions in artificial forest edge meadows and open forest floors, might promote a rapid reduction in the populations of such dwarf plants.