The fruit morphometric variation and fruit type evolution of the stone oaks (Fagaceae, Lithocarpus)

The great species diversity of Lithocarpus is associated with interspecific fruit morphological variation, represented by acorn (AC) and enclosed receptacle (ER) fruit types. Species representing both fruit types co-occur in the same forests and share two distribution centers in southern China and southeastern Asia. The predation selection hypothesis suggests that the fruit morphological mechanical trade-off between two fruit types could represent divergent dispersal strategies under varied predation pressures. By integrating phylogenetic construction with fruit morphometric study, we tried to verify the predation selection hypothesis and elucidate the fruit type evolution of Lithocarpus, which is critical in interpreting the distribution and diversification of the genus.

We identified the functional trade-off between the two fruit types: ER species have bigger seeds which are enclosed mainly by receptacle representing stronger physical defense; whereas the seeds of AC species are smaller and enclosed mainly by thin pericarp representing lower mechanical protection. Despite some reversals from ER back to AC, the ancestral state reconstruction in combination with thermal analysis supports the hypothesis that ER is the derived fruit type from AC-like ancestors independently across all clades.

Our results support the predation selection hypothesis by verifying the mechanical trade-off between the two fruit types. We propose a divergent selection theory for the two fruit types: the seed size and mechanical defense of AC species become smaller, whereas those of ER species become larger and require more morphological modifications in the receptacle. This signified the importance of the receptacle in differentiating the two fruit types and in the fruit morphological modification through evolutionary time. We found that the ER-type species evolved independently in all clades and across varied climates from tropical to warm temperate regions. As ER fruits are products of convergent evolution, we propose to examine the predation and dispersal variation between two fruit types in the future to verify whether predation selection is the reason behind fruit type evolution of the stone oaks.