“…Floral visitors usually follow an optimal and possibly consistent route within inflorescences, which results in the maximum net rate of food gain (Heinrich, 1975;Hainsworth et al, 1983;Jordan & Harder, 2006). Plants possibly minimize occurrences of geitonogamy according to their visitors' behavior by spatial and temporal arrangement of flowers (Iwata et al, 2012;Jordan et al, 2016). Bee pollinators usually tend to start foraging at lower flowers and move upward in vertical inflorescences (Darwin, 1877;Utelli & Roy, 2000;Fisogni et al, 2011;Zhao et al, 2016); the placement of older and thus female flowers below younger male flowers in many protandrous plants is thought to have evolved to reduce geitonogamy, because lower female-phase flowers would receive pollen from other plants whereas upper male-phase flowers would disperse pollen to other plants (Harder et al, 2000;Routley & Husband, 2003;Jers akov a & Johnson, 2007;Liao & Harder, 2014).…”