Calceolariaceae: floral development and systematic implications.

The recent establishment of the new family Calceolariaceae, separated from Scrophulariaceae on the basis of molecular evidence, is complemented here by a scanning electron microscopy study of floral morphology and development of 12 species encompassing all genera (Calceolaria, Jovellana, and Stemotria [= Porodittia]). All species showed a similar pattern of organ initiation. The slightly zygomorphic, four-merous calyx is the first floral organ series initiated, with the primordia emerging consecutively in a unidirectional (dorso-ventral) succession. The two entire corolla lips in Calceolaria and Jovellana arise as uniform meristematic ridges (sometimes with a central emargination, especially in Jovellana), kept apart by two lateral stamen primordia. Later the margins of the lips fuse across the backs of the young stamens, giving rise to the short corolla tube (late sympetaly). Stemotria stands out by having three stamens instead of two and a bilobed lower lip, resulting in a trimerous corolla. Similar architecture was found in teratological flowers of Calceolaria. The perianth of Calceolariaceae is shown to be derived from a tetramerous condition, not from pentamery as traditionally believed. This is in agreement with the separation of Calceolariaceae from Scrophulariaceae and with their placement in succession of Oleaceae and Tetrachondraceae in the basal Lamiales. The hitherto puzzling molecular evidence is thus supported by morphological-developmental features of the flower.