Phenological and morphological variations of Oryza rufipogon and O. nivara in Sri Lanka and their evolutionary implications.

Phenological and morphological variation are widely viewed as a pivotal driver of ecological adaptation and speciation. Here, we investigate variation patterns of flowering phenology and morphological traits within and between O. rufipogon and O. nivara populations in Sri Lanka by incorporating the in situ observation in natural habitats and manipulative experiments in the common gardens. First, we observed varying degrees of phenological variation under different temporal and spatial conditions, suggesting that flowering phenology of two Oryza species varied depending on both environments and management practices. Particularly, the Sri Lankan O. nivara exhibits high plasticity in flowering phenology, implying that O. nivara might not be an annual in the strict sense. Second, the observation that flowering time of the two species overlapped suggests that the primary factor to maintain the species divergence in Sri Lanka may not be flowering time but rather environments. Third, our selection analysis suggests that interspecific divergence in the traits related to reproduction and habitat preference is adaptive and most likely driven by natural selection. Together, our case study on the Sri Lankan O. rufipogon and O. nivara enhances the understanding of the roles of phenotypic plasticity and environmental factors in the processes of adaptation and speciation.