Associated morphometric and geospatial differentiation among 98 species of stone oaks (Lithocarpus)

Two fruit types can be distinguished among stone oaks (Lithocarpus) species: the ‘acorn’ (AC) and the ‘enclosed receptacle’ (ER) types. Our morphometric analysis of 595 nuts from 98 species (one third of all Lithocarpus spp.) found substantial transition in mechanical protection of the seed between two woody fruit tissues (exocarp and receptacle) of two fruit types. AC fruits were smaller in seed and fruit size and the thin brittle exocarp largely enclosed the seed, whereas ER fruits were larger and the seed was mostly enclosed by thick woody receptacle tissue. The differences in these two tissues were considerably greater between compared to within fruit type and species. Geospatial distribution showed that seed size of all examined species increased with elevation and decreased with latitude, the physical defense increased with both elevation and latitude, and ER-fruit species were more common at higher elevation. The two fruit types represent distinct suites of associated traits that respond differently to the various biotic and abiotic factors associated with geographic variation, profoundly impacting the evolution of the two fruit types. The co-occurrence of two fruit types in the same forest could be a consequence of distinct fruit and animal interactions.