CONE DEVELOPMENT IN LIBOCEDRUS (CUPRESSACEAE)—PHENOLOGICAL AND MORPHOLOGICAL ASPECTS

Abstract

Cones in Libocedrus plumosa are initiated in New Zealand in February (late summer) and continue to develop through winter. The ultimate pair of bracts protrude and continue extension growth while the apex remains flat. Ovules are initiated in two pairs in July on the cone apex (i.e., are axial in origin), alternate with the upper two pairs of bracts. Ovule differentiation proceeds rapidly to the stage of pollination; the cone apex may develop further as a short columella. Soon after ovule initiation an adaxial ligulelike outgrowth of each member of the upper two pairs of bracts is initiated, that of the ultimate pair being broader than that of the lower pair. By intercalary growth of each ligule base an enlarged structure is developed, displacing the bract into a lateral position and forming the four scales that enclose the developing seeds in a valvate manner. Cone and seed maturation follows with seed dispersal in March, approximately 1 year after cone initiation. The results show that there is no morphologically discrete ovuliferous scale, and there is no ontogenetic fusion of discrete separate structures. The vascular system of the scale complex develops after the ovules are initiated and forms a single series of vascular bundles with inversely oriented xylem and phloem; this and other histological changes are the result of the activity of the intercalary growth process and do not relate to primary structures. The results are discussed in relation to existing knowledge of cone development in Cupressaceae, in which axially borne ovules are common.