Flowers and floral biology of the holy grasses (Hierochloe and Anthoxanthum: Aveneae, Gramineae)

Trifloreted spikelets in species of Anthoxanthum and of Hierochloe are united by the syndrome of two lower male or neuter florets and a differentiated third floret, perfect, bistaminate, and protogynous. Differences in presentation of the floral organs from the third anthoecium separate the genera. Anthers in Anthoxanthum, longer than the f3 anthoecium, keep it open until they migrate on elongating filaments, following behind the protogynous stigma-styles. Both emerge near the apex of the upper glume which tightly encloses all three florets. In Hierochloe the uppermost anthoecium is closed over the androecium and gynoecium until lodicule-controlled anthesis when stigma-styles are exserted unhindered by glumes, and the anthers are exserted later on long filaments from the chasmogamous lower florets. In Hierochloe triandrous lower florets are constant. In Anthoxanthum the two lower florets are neuter in Eurasian species, but in species in Africa, Asia, and Malesia, sometimes f1 florets are also staminate, and very exceptionally both florets present stamens. These reflect a sexual variability in Anthoxanthum unknown in Hierochloe. Data from flowers of 30 species of Hierochloe and 17 species of Anthoxanthum are tabulated; included are ratios of f3 anthers:anthoecium and stigma-styles:the upper glume, G2, two ratios which most reliably reflect the intergeneric floral patterns. Pathways to those varied floral systems are outlined. A sectional taxonomy is proposed for Hierochloe: Sect. Hierochloe autonymum; Sect. nova Monoecia for species in South America where monoecism occurs in all taxa. Andromonoecism is the predominant sexual system in boreal and austral species of Hierochloe, but is interrupted by apomixis in northern Europe.