Structure and development of the ovule and seed in Aristolochiaceae, with particular reference to Saruma

All members of Aristolochiaceae have anatropous, bitegmic, crassinucellate ovules, which are endostomic except in Saruma and Asarum arifolium where ovules are amphistomic. The outer integument is two cell-layered and the inner integument is three cell-layered. The chalazal megaspore is the functional one. All these conditions appear to be plesiomorphic for the order Piperales, which consists of five families, Aristolochiaceae, Hydnoraceae, Lactoridaceae, Piperaceae and Saururaceae. The embryo sac in Aristolochiaceae is eight-nucleate and corresponds to the Polygonum type; a hypostase is frequently present in this family. The seed coat of Aristolochia s.l., Asarum, Saruma and some Thottea species consists primarily of a two cell-layered testa, and a three cell-layered tegmen. In some species the cells of the outer epidermis become radially elongated, forming reticulate wall thickenings. Cells of the inner layer of the testa have crystals and thickened inner walls. The three layers of the tegmen are tangentially elongated, and become cross fibres at maturity, as fibres of the outer and inner layers are parallel to the seed axis, whereas those of the middle layer are perpendicular to it. This type of seed coat anatomy is synapomorphic for Aristolochiaceae. In addition, the gross morphology of the seed and elaiosome histology are remarkably similar in Asarum and Saruma, thus supporting a sister-group relationship between them. Embryological and seed characters do not supply any synapomorphy that support a close relationship between Aristolochiaceae, Hydnoraceae and Lactoridaceae. Instead, some seed features such as the absence of seed appendages and the collapsed cells of endotesta may indicate a close relationship of Lactoris with Piperaceae plus Saururaceae, although this is the subject of further analysis.