Fleshy Structures Associated with Ovule Protection and Seed Dispersal in Gymnosperms: A Systematic and Evolutionary Overview

Fleshy structures associated with the ovule/seed arose independently several times during gymnosperm evolution. Fleshy structures are linked to ovule/seed protection and dispersal, and are present in all the four lineages of extant gymnosperms. The ontogenetic origin of the fleshy structures could be different, and spans from the ovule funiculus in the Taxus baccata aril, the ovule integument in Ginkgo biloba, to modified bracts as in case of Ephedra species. This variability in ontogeny is reflected in the morphology and characteristics that these tissues display among the different species. This review aims to provide a complete overview of these ovule/seed-associated fleshy structures in living gymnosperms, reporting detailed descriptions for every genus. The evolution of these independently evolved structures is still unclear, and different hypotheses have been presented—protection for the seeds, protection to desiccation—each plausible but no one able to account for all their independent origins. Our purpose is to offer an extensive discussion on these fleshy structures, under different points of view (morphology, evolution, gene involvement), to stimulate further studies on their origin and evolution on both ecological and molecular levels.