Floral morphology and embryology of Helwingia (Helwingiaceae, Aquifoliales): systematic and evolutionary implications

Helwingia, a shrub or small tree of four species distributed in East Asia, has been assigned to various families, mainly Cornaceae. However, molecular analyses show that the genus belongs to its own family Helwingiaceae which is sister to Phyllonomaceae (Phyllonoma only) in the order Aquifoliales. On the basis of H. japonica, we investigated the poorly understood floral and embryological characters of Helwingia, and compared their features with those of other Aquifoliales, particularly those of Phyllonomaceae. Results showed that perianth leaves of Helwingia represent sepals, because in plesiomorphic pentamerous flowers, they agreed in position with sepals (not with petals) in pentamerous flowers of Phyllonoma. Overall comparisons based on available information show that, while sharing with Phyllonoma the epiphyllous inflorescence, the inferior ovary, and an epigynous disc nectary as syapomorphies, Helwingia is characterized by loss of petals, obhaplostemony, large recurved stigmas, poorly developed disc nectary, tenuinucellate ovules with a mature female gametophyte filled with densely stained cytoplasm, and a thin mature seed coat. Morphological evidence, like molecular evidence, confirms that Helwingia is sufficiently distinct to be placed in its own family. Morphological and field observations suggest wind and insect pollination in H. japonica, which is the first example of ambophily in Aquifoliales.