Developmental anatomy of seedlings of Indodalzellia gracilis (Podostemaceae)

In Tristichoideae, aquatic angiosperms in the family Podostemaceae, Terniopsis, Tristicha, Indotristicha and Cussetia have creeping roots with flanking (sub)cylindrical shoots, while Dalzellia is rootless and has crustose shoots. Indodalzellia gracilis, sister to a clade of Dalzellia zeylanica and Indotristicha ramosissima, has subcrustose shoots on the side of creeping roots, suggesting that I. gracilis may be a key species to reveal how saltational evolution of the body plan occurred in these three species. We investigated developmental morphology of I. gracilis seedlings grown in culture, using scanning electron microscopy and semi-thin serial sections. As in D. zeylanica, the plumular apical meristem in the seedling gives rise to two shoot apical meristems, which develop into horizontal subcrustose shoots with dorsal and marginal leaves. Neither radicle nor adventitious root is produced from the hypocotyl, but an adventitious root arises endogenously from the juvenile shoot and from some shoots of adult plants. These results, together with the phylogenetic relationships, suggest that the Indodalzellia seedling evolved by loss of the adventitious root derived from the hypocotyl, appearance of shoots in the axil of cotyledons, and appearance of adventitious roots from adventitious shoots. The difference in place of origin of the root between Indodalzellia and I. ramosissima suggests differing evolutionary origin of the root in Tristichoideae.