Morphological and ontogenetic studies on southern African podocarps. Initiation of the seed scale complex and early development of integument, nucellus and epimatium

The primordium of the seed scale complex (ssc) (ovule with epimatium) is initiated in the axil of the first or second cone bracts (prophylls) as a dome shaped structure consisting of a group of uniform, meristematic cells. A distinct protodermal layer develops. The nucellus is a dome on the ventral side of the ssc primordium. In the species of section Podocarpus the integument is initiated as a circular ridge around the nucellus, while in P.falcatus two protrusions on the anterior and posterior sides of the nucellus are the first indications of integumentary differentiation. The integument of all species studied is of subdermal origin. The epimatium (sensu stricto) is initiated after the integument, is of subdermal origin and forms a hood around the developing ovule. Considering research results, together with related literature, it is concluded that the integument of gymnosperms may be homologous with the outer integument of a bitegmic angiospermous ovule, that the position of integumentary initiation may be specific for certain taxa and that there seems to be no constant sequence of emergence of ovular envelopes in gymnosperms.