Developmental anatomy of Terniopsis malayana (Podostemaceae, subfamily Tristichoideae), with implications for body plan evolution

The developmental morphology of Terniopsis malayana, an unusual aquatic angiosperm from Thailand, was examined. A unique vegetative structure called the ramulus arises endogenously in the root tissue. The ramulus has an actively growing apical meristem. The ramulus branches several times to form a ramulus complex consisting of up to six ramuli, which are distichously arranged in almost a single plane. In a ramulus complex, the new ramulus (ramulus branch) is initiated on the adaxial side of the first (the basalmost) scale in the first ramulus, but at a site lateral to the first scale in later ramuli, suggesting that the new ramulus arises from axillary or extra-axillary buds of the immediately older ramulus. Ramulus growth is terminated in association with the loss of the apical meristem, and its axillary or extra-axillary buds begin to grow to form the next new ramulus instead. The flower occurs in place of the youngest ramulus, when reproductive. It seems likely that the Terniopsis ramulus and its scale are comparable to the shoot and leaf, and thus a ramulus complex is interpreted as a sympodially branched shoot.