Floral Ontogeny of Zippelia begoniaefolia and its Familial Affinity: Saururaceae or Piperaceae?

Abstract

Zippelia begoniaefolia Bl., a monotypic species having characteristics of both Piperaceae and Saururaceae, has racemes of about 20 small flowers lacking a perianth, each with six free stamens and a four-carpellate syncarpous gynoecium. The inflorescence apical meristem initiates bracts acropetally and helically, each of which subtends a later initiated single floral apex; there are no “common” primordia. The six stamens are initiated as two lateral pairs and two solitary successive primordia, the latter two opposite in median sagittal positions. Four carpel primordia are initiated as a lateral pair and two successively initiated in the median sagittal plane. This order of organ inception is unique among Piperaceae and Saururaceae. Intercalary growth below carpellary attachment raises them up on a common cylindrical base that becomes the syncarpous ovary, covered with unique glochidiate hairs and containing a single basal ovule. The free portions of the carpels become the reflexed papillate stigmas. The floral vascular system has a single bundle at base that branches to supply the bract and flower traces. The floral vasculature is similar but not identical to that of Saururus (Saururaceae) and some Piper species (Piperaceae). Plesiomorphic character states of Zippelia that are shared with Saururus include hypogyny, free stamens, cleft stigma, and a similar floral groundplan. Synapomorphies, derived shared character states that unite Zippelia with Piperaceae, include syncarpy, solitary ovule, basal placentation, fused ventral carpellary bundles, and a double vascular cylinder in the stem. Cladistic analysis aligns Zippelia with Piperaceae because they share apomorphies, and because Zippelia shares only plesiomorphies with Saururus.