Seed coat morphology and its systematic implications in Cyanea and other genera of Lobelioideae (Campanulaceae)

Recent surveys of seed coat morphology in Lobelioideae (Campanulaceae) have demonstrated the systematic utility of such data in the subfamily and led to a revision of the supraspecific classification of Lobelia. Expanding upon these studies, we examined via scanning electron microscopy 41 seed accessions, emphasizing lobelioid genera in which only one or no species had been examined. Most conformed to previously described testal patterns. However, five species of the endemic Hawaiian genus Cyanea, comprising the molecularly defined Hardyi Clade, had a unique testal pattern (here termed Type F), characterized by laterally compressed, almost linear, areoles with rounded, knob-like protuberances on the radial walls at opposite ends. This offered a convenient synapomorphy for recognition of a clade originally defined on a molecular basis. A second unique testal pattern was found in the related Hawaiian endemics Brighamia and Delissea, thus supporting their close relationship. In this type (here termed Type G), the seed coat is irregularly wrinkled (rugose), creating broad, rounded ridges that run more-or-less perpendicular to the long axis of the seed and thus to the long axis of the testal cells. Seed coat morphology also supported the monophyly of all 124 species of Hawaiian Lobelioideae and their probable derivation from Asian species of Lobelia subg. Tupa. Additional studies supported close relationships between (1) the neotropical genera Centropogon and Siphocampylus; (2) the western American genera Legenere and Downingia; and (3) Jamaican Hippobroma and Lobelia sect. Tylomium, a group endemic to the West Indies.