Comparative floral structure and systematics in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae)

Abstract

Floral morphology, anatomy and histology were studied in representatives of all families of current Oxalidales, which were recently constituted as a result of molecular systematic studies by other authors, and are composed of families of different positions in traditional classifications (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Two of the three pairs of sister (or nested) families that come out in molecular analyses are highly supported by floral structure: Oxalidaceae/Connaraceae and Elaeocarpaceae/Tremandraceae, whereas Cephalotaceae/Cunoniaceae are not especially similar at the level of Oxalidales. Oxalidaceae and Connaraceae share petals that are postgenitally united into a basal tube (although they are imbricate in both) but free at the insertion zone, stamens that are congenitally united at the base, uniseriate glandular hairs on the stamen filaments, and ovules that are hemianatropous to almost orthotropous. The sharing of a special type of sieve-tube plastids and of trimorphic heterostyly, studied by other authors, should also be mentioned. With Brunelliaceae, the two families share an androgynophore and nectaries at the base of the stamens in alternisepalous sectors. Elaeocarpaceae and Tremandraceae share buzz-pollinated flowers and a syndrome of features functionally connected with it. In addition, petals are larger than sepals in advanced bud, they are valvate, involute and enwrap part of the adjacent stamens, they have three vascular traces. Lignified hairs are common on the anthers and are found in the ovary locules and on the ovules (not lignified) of representatives of both families. Ovules have a chalazal appendage, and the inner integument is much thicker than the outer.