The roles of the ostiole in the fig-fig wasp mutualism from a morpho-anatomical perspective

The syconium is the urn-shaped inflorescence shared by all species of the genus Ficus. The orifice at the apex of the syconium is called the ostiole, and it is covered by interlocking bracts. The ostiolar bracts can have different arrangements, which only allow the entry of mutualist wasps and promote reproductive isolation among Ficus species. Here, we analyze the ostiolar structures that could play a role as selective filter and therefore impact the fig-fig wasp mutualism in the neotropical Ficus sections Americanae and Pharmacosycea. Samples of syconia with pistillate flowers during the receptive phase of seven species of Ficus were examined using light and scanning electron microscopy. Tests for histolocalization of substances were employed to detect secretory activity throughout the ostiolar tissues. Our results indicated that the ostiole has two components: ostiolar bracts and the periostiolar zone. Interspecies variation in ostiolar bract arrangement in both sections studied was broader than previously reported. We report for the first time for Ficus: (i) two types of ostiolar osmophores (mesophyll and diffuse), that could be a source of volatile compounds for attracting fig wasps; (ii) colleters in the axil of ostiolar bracts, which probably lubricate and facilitate the entry of pollinating wasps into the syconial cavity; (iii) secretory trichomes around the ostiolar bracts, and (iv) syconium basal bracts (F. isophlebia) covering the ostiole, which are the first physical barrier that the fig wasps must overcome to access receptive pistillate flowers. We describe the zones that compose the ostiole, which support the hypothesis that the ostiole is a selective filter in the interactions of fig trees with Agaonidae fig wasps. We also suggest that ostiolar osmophores, colleters, the periostiolar zone, and the arrangements of the ostiolar bracts may be informative with respect to Ficus systematics.