Developmental features of the discontinuous stem vascular system in the rattan palmCalamus(Arecaceae‐Calamoideae‐Calamineae)

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Abstract
Calamus is a climbing palm marked by considerable internodal extension and limited stem-thickening growth, but with a surprisingly discontinuous axial vascular system. Stem bundles end blindly in a basipetal direction and are connected to each other only by narrow and late-developing transverse commissures. Vascular connection via leaf traces between stem and leaf is made over about nine plastochrons (P), but the dominant central system is completed by about P7, with subsequent bundles forming the crowded fibrous peripheral system, which has reduced or no vascular tissues. The stem internode below a leaf completes its extension and maturation only by P10 to P11. Axial stem bundles originate as procambial strands that are discontinuous apically for up to 15 plastochrons before being “captured” by a developing leaf. Their distal unconnected ends arise by dedifferentiation of ground parenchyma cells. Protoxylem is initiated as short overlapping initials that differentiate progressively during extension growth, which ruptures all but the last-formed elements. Their form, with tapered ends, means that they mature as tracheids. Metaxylem appears only late in shoot development, shortly before internodal elongation ceases (P8) and always unconnected to the late-differentiating protoxylem. In each axial bundle protophloem differentiates as a single strand, subsequently and much later appearing as two separate metaphloem strands as the early initials, ruptured by extension growth, are replaced by fibers. It is suggested that the unique features of this stem can be ascribed to the absence of a “meristematic cap,” which otherwise typifies palms of normal habit, and that discontinuity is causally related to the pronounced late stem extension growth.