Reproductive phenological shifts and other phylogenetic trait changes in the Arbutoideae (Ericaceae) in the context of drought, seed predation, and fire

Phenology is an ecologically critical attribute that is commonly coordinated with other plant traits. Phenological shifts may be the result of evolutionary adjustments to persistently new conditions, or they may be transitory, varying with annual fluxes in abiotic conditions. In summer-dry, fire-prone Mediterranean-type climates, for example, many plant lineages have historically migrated from forests to more arid shrublands resulting in adaptive trait changes. These shifts in habitat abiotic conditions and biotic interactions influence morphology of flowers and fruits and interact with phenological timing. The Arbutoideae (Ericaceae) is one lineage that illustrates such modifications, with fruit characters evolving among genera from fleshy to dry fruit, thin to stony endocarps, and bird to rodent dispersal, among other changes. We scored herbarium collections and used ancestral trait analyses to determine phenological shifts among the five Arbutoid genera found in semi-arid climates. Our objective was to determine if phenology shifts with the phylogenetic transition to different reproductive characters. Our results indicate that phenological shifts began with some traits, like the development of a stony endocarp or dry fruits, but not with all significant trait changes. We conclude that early phenological shifts correlating with some reproductive traits were permissive for the transition to other later character changes.