Morphology and morphogenesis of ensiform leaves, syndesmy of shoots and an understanding of the thalloid plant body in species of Apinagia, Mourera and Marathrum (Podostemaceae)

Abstract

The flattened, irregularly shaped and lobed or dissected leaves of Apinagia riedelii, A. latifolia, A. goejei, Mourera aspera and Marathrum utile (subfamily Podostemoideae) are ensiform in structure. After the typical bifacial inception, further growth of leaves proceeds in the midrib area, i.e. in the median plane of the leaf. The lower leaf zone is characterized by a sheath that orientates ‘at the side of the blade’, i.e. at the adaxial edge of the sword-like leaf. The ensiform blades are lobed, incised or pinnately dissected with the tips terminated in elongated threads or thin filaments. Leaves of Apinagia riedelii resemble pinnately dissected compound leaves. The dissected structure represents a secondary superimposition of the ensiform shape and a parallel development to compound leaves. It is interpreted as an adaptation to the rapid current, established in the floating shoots of these aquatic plants. The basal portions of main shoots and successive branches are fused due to a retarded separation. Fusion of this kind has been termed a syndesmy. The fused region of shoots is superseded by the fusion of the lower leaf zones of (distichously positioned) adjacent leaves, occurring at their margins. The fused leaf bases form a cavity for the terminal flower bud of each shoot and cover it in the form of a hood. The flower buds are hidden from external view. The meristematic growing zones are thus protected and enclosed within the syndesmic plant body which, in this way, attains the ‘thalloid’ appearance especially developed in Apinagia goejei and A. latifolia but also present in the other species. The results of this study enable an understanding of the particular appearance of these Podostemoideae as modifications of the typical structures according to the ‘principle of variable proportions’.