The effect of seasonality on developmental stages of anthetic ovule integuments in Mediterranean orchids

Orchids, differently from most flowering plants, have under-differentiated ovules at anthesis that require pollination to complete differentiation. This ovule developmental stage has been often observed in tropical species in which the absence of an evident seasonality may allow plants to extend their phenology beneficiating of a long time for post-pollination events. Here, we used scanning electron microscopy (SEM) to detect ovule integument developmental stages in 21 species of Mediterranean Orchidoideae and Epidendroideae and in 11 tropical Epidendroideae with the aim of understanding whether species with a seasonal constraint and shorter time for post-pollination ovule maturation are characterized by different stages of ovule development at anthesis. We found that Mediterranean orchids (both Orchidoideae and Epidendroideae) have more developed ovule integuments than tropical orchids. Most species show partially to fully developed ovules at anthesis with the exception of Cephalanthera where ovules are arrested in a very early developmental stage. Despite the phylogenetic signal, anthetic ovule integument development differs between related species (with different pollination strategies or blooming times), suggesting the presence of some ecological constraints. The synchronization between ovule integuments and megagametophyte development, as found in tropical orchids, is also confirmed in Mediterranean orchids. Our data show that Mediterranean and tropical orchids clearly differ in anthetic ovule developmental stages, likely depending on seasonality.