The systematic relevance of fruit and seed structure in Bersama and Melianthus (Melianthaceae)

Abstract.

 The fruit and seed anatomy and morphology of the two genera Bersama and Melianthus (Melianthaceae, Sapindales) have been studied in an effort to clarify their systematic position. On the basis of the differences in pericarp and seed anatomy as well as in other exomorphic characters the segregation of Bersama into a distinct family Bersamaceae is supported. Evidence mainly from seed anatomy and morphology emphasizes the anomaly of the traditional inclusion of Bersama and Melianthus in the Sapindales, since they have a distinct seed-coat structure and seed vascularization. The fruit and seed anatomy does not confirm any relationships with alternatively suggested exo-mesotestal Lardizabalaceae. The exotestal seed coats of Bersama and Melianthus with a differentiated palisade of Malpighian cells in the exotesta, dimerous raphal vascular skeleton, abundant endosperm, and a small differentiated straight embryo show a resemblance with the exotestal albuminous seeds of Rhamnaceae and Elaeagnaceae. Using also additional data on fruit, floral and vegetative morphology it is suggested that Bersamaceae with Melianthaceae and Rhamnaceae/Elaeagnaceae constitute a distinct relict side-branch of exo-mesotestal rosidaceous ancestry, the new order Melianthales in the superorder Rhamnanae (Rosidae). The formerly suggested relationship of this side-branch to exotegmic Malvales is not supported by seed anatomy. The affinity with exotegmic Celastrales, which are considered as a possible connecting link between archaic exo-mesotestal Rosales and exotestal Rhamnales/Elaeagnales, is also found untenable.