Functional and ecological diversification of underground organs in .

The evolution of geophytes in response to different environmental stressors is poorly understood largely due to the great morphological variation in underground plant organs, which includes species with rhizomatous structures or underground storage organs (USOs). Here we compare the evolution and ecological niche patterns of different geophytic organs in L., classified based on a functional definition and using a clade-based approach with an expert-verified specimen occurrence dataset. Results from PERMANOVA and Phylogenetic ANOVAs indicate that geophytic species occupy drier areas, with rhizomatous species found in the hottest areas whereas species with USOs are restricted to cooler areas in the montane tropics. In addition, rhizomatous species appear to be adapted to fire-driven disturbance, in contrast to species with USOs that appear to be adapted to prolonged climatic disturbance such as unfavorable growing conditions due to drought and cold. We also show that the evolution of rhizome-like structures leads to changes in the relationship between range size and niche breadth. Ancestral state reconstruction shows that in rhizomatous species are evolutionarily more labile compared to species with USOs. Our results suggest that underground organs enable plants to shift their niches towards distinct extreme environmental conditions and have different evolutionary constraints.