Detailed seed cone morpho-anatomy of the Prumnopityoid clade: an insight into the origin and evolution of Podocarpaceae seed cones.

Abstract

Seed cone traits are significant for understanding the evolutionary history of conifers. Podocarpaceae has fleshy cones with a distinct morphology compared with other conifers. However, we have a poor understanding of the seed cone morphology of the Prumnopityoid clade and within Podocarpaceae. This study presents detailed seed cone morpho-anatomy and the evolution of fleshy structures traits in the Prumnopityoid clade. We investigated the detailed seed cone morpho-anatomy of selected species from the nine genera using the histological method. The evolution of morpho-anatomical traits was assessed using ancestral state reconstruction methods. The Prumnopityoid clade has evolved fleshy seed cones using different functional structures (e.g. aril, epimatium, bracts or receptaculum) and fleshiness is an ancestral trait in the clade. An epimatium is present in all genera except Phyllocladus, but with different structural morphologies (e.g. a fleshy asymmetrical cup-like epimatium or an epimatium that is fused with the integument, forming a fleshy sarcotesta-like seed coat). In all species with fleshy sarcotesta-like seed coats, the endotesta is hard and woody, forming a sclerotesta-like structure and the epimatium and exotesta are fused, forming a fleshy sarcotesta-like structure. This study highlights that the Prumnopityoid clade has an amazing diversity of structures and complex evolutionary patterns. Fleshiness is an ancestral trait of the clade and has been achieved via diverse evolutionary pathways and structures. This clade has four distinct seed cone types, i.e. drupe-like, receptaculate, arilloid and dacrydioid cones, based on morpho-anatomical structures and traits. The macrofossil record also demonstrates the presence of several structures and traits.