Pollen and reproductive morphology of Rhigiophyllum and Siphocodo (Campanulaceae): two unique genera of the fynbos vegetation of South Africa

Abstract

Pollen grains of Rhigiophyllum squarrosum Hochst., Siphocodon spartioides Turcz. and S. debilis Schltr., are flattened and triangular with pores at the angles. This morphology is radically different from known pollen of the Campanulaceae s.sfr:: the Campanulaceae are treated here as a family separate from the Lobeliaceae, Cyphiaceae, Nemacladaceae, Pentaphragmataceae and Sphenocleaceae (Lammers 1992). As traditionally conceived, the Campanulaceae is very heterogeneous and, in many classifications, these families were treated as subfamilies of a much-enlarged Campanulaceae. The consistently different floral morphology, biochemistry and pollen structure of the Lobeliaceae favours the recognition of this predominantly tropical group as a separate family.The pollen grains of these species are described in comparison with other members of the Campanulaceae. Based on surface characteristics of their pollen grains, we conclude that they represent an early offshoot o f the wahlenbergioid line­age in southern Africa. We suggest that this unique pollen may also be the result of a highly selective regime in the fynbos, associated with specialized pollinators, and base-poor soils, in addition to possible adaptations for ant dispersal and fire. Rhigiophyllum Hochst. and Siphocodon Turcz. are also unique in having free carpel-like structures within the ovary. These shrink to form seed pockets around the seeds and disperse as units when the capsule matures. Data from molecular studies support the contention that these taxa form a sister group to all other wahlenbergioids and that this should be formally recognized in a classification system. We treat Rhigiophyllum and Siphocodon within the Campanulaceae: Wahlenbergioideae, as a separate tribe, the Rhigiophylleae tribus nov., the species of which are distinguishable from other wahlenbergioids by unique angulaperturatc pollen, epipetalous stamens, free carpel-like structures and seed pockets.