Confirmed dioecy in at least 15 solanums is distributed across the genus, with perhaps 6 independent origins, and with crypticity in the form of morphologically hermaphroditic, but functionally unisexual, flowers characterizing all species. Dioecy is not more strongly associated with islands. Inaperturate pollen in pistillate flowers characterizes almost all, but not the two dioecious species studied herein. Dioecy in both species indicates leakiness (rare hermaphroditic flowers on male plants) that helps explain island colonization and radiation. Leakiness allowed confirmation-usually impossible for dioecious species-of self-compatibility for S. polygamum, and thus support for the hypothesis that dioecy evolved to promote outcrossing.
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We confirmed functional dioecy of Withania aristata via field and greenhouse studies. Male flowers are significantly larger. Female flowers bear stamens with no pollen; males bear 220 000 grains. Stigmata of male flowers senesce in buds. Anatomical observations confirm more ovules in females and an ovarian nectary in both sexes. We detected nectar in female flowers in the greenhouse but found no nectar in males. Thus, males offer pollen and females nectar. Females bear large numbers of fruits and, infrequently, male plants bear few significantly smaller fruits with few seeds. Outcrosses of females (self crosses impossible without pollen) yielded fruits in young buds, older buds, and open flowers. Self crosses of male flowers succeeded only with very young buds. Although functionally dioecious, this species manifests self-compatibility; however, no fruits are produced autonomously. Bee species (Lassioglossum, Amegilla, Apis) visit flowers and mature buds. Bud visits in which bees force petal tips apart, coupled with self-compatibility, may explain infrequent fruit on males. Thus, dioecy in W. aristata seems to have evolved from self-compatible ancestors, that leaky dioecy may have been favored during colonization, and, that despite autogamy and a low floral visition rate, this endemic enjoys a high rate of reproductive success.
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