Specialized structures in the leaf epidermis of basal angiosperms: morphology, distribution, and homology

The morphology of specialized structures in the leaf epidermis of 32 species of basal (ANITA: Amborella, Nymphaeales, Illiciales, Trimeniaceae, and Austrobaileyaceae) angiosperms, representing all seven families and 11 of 14 genera, was investigated using light and scanning electron microscopy. Distribution, density, and size of structures were also measured, and character evolution was analyzed. Hydropotes are a synapomorphy of Nymphaeales and ethereal oil cells are a synapomorphy of Austrobaileyales, but uniseriate nonglandular trichomes appear to have arisen independently several times. Specialized structures are frequently characterized by adjacent epidermal cells that have striking similarities in their form and arrangement (i.e., architecture) to subsidiary cells of certain types of stomatal complexes. Additionally, forms intermediate to oil cells and stomata, to trichomes and stomata, and to hydropotes and oil cells are present in some taxa. Thus, all of these specialized structures and their adjacent epidermal cells form complexes that may be homologous with, and evolutionarily derived from stomatal complexes, and the specialized structure, or portion thereof, may be homologous to the stoma or guard mother cell. Improved knowledge of the morphology and evolution of these structures in the earliest branching extant angiosperm lineages has a bearing on many diverse areas of botany.