Micromorphology and systematic distribution of pit membrane thickenings in Oleaceae: Tori and pseudo-tori

Recent studies on the functional significance of pit membranes in water conducting cells have renewed general interest in their micromorphology. At least two types of pit membrane thickenings have been described in angiosperm families, i.e. genuine tori and pseudo-tori. This study explores the distribution and morphology of pit membrane thickenings in 69 species and 23 genera within Oleaceae using light and electron microscopy. Torus-bearing pit membranes are confirmed for Osmanthus, and new records are reported for Chionanthus retusa, Picconia azorica, and P. excelsa, but not for the other species studied of Chionanthus. This infrageneric variation suggests that tori represent a plastic feature that has evolved more than once within the family as the result of functional adaptation to efficient and safe water transport. Pseudo-tori are observed in species of Abeliophyllum, Fontanesia, Forsythia, Jasminum, Ligustrum, Menodora, and Syringa. Based on structural differences, we state that tori and pseudo-tori can be distinguished as non-homologous features.