Environmental influence on primary sex ratio in a dioecious plant

Abstract
The proximity of mates can influence mating opportunities and the quantity and quality of offspring, especially in dioecious plant species. Progeny sex ratios modulated by environmental conditions is one of the most radical ways in which offspring quality may be influenced, yet it has rarely been reported in plants. A mechanism proposed to influence progeny sex ratios in dioecious plants involves competition between female- and male-determining microgametophytes (certation) as a result of variation in pollination intensity. However, the role of selective fertilization in dioecious plants is controversial and has not been demonstrated under field conditions. Here we investigate whether natural variation in the spatial arrangement of females and males influences pollination intensity and progeny sex ratios in the wind-pollinated herb Rumex nivalis. Based on previous experimental manipulation of pollination intensity in this species, we predicted that maternal parents in close proximity to males would produce more strongly female-biased progeny sex ratios. We tested this prediction in six alpine populations in Switzerland by measuring the distance between focal females and neighboring males and assessing pollen loads and seed sex ratios of maternal parents. In four of the six populations, females positioned in close proximity to males captured more pollen and exhibited more female-biased sex ratios. Our results demonstrate that demographic aspects of the maternal mating environment can influence progeny sex ratios. The most probable explanation for biased primary sex ratios in Rumex is selective fertilization resulting from pollen tube competition.