Studies on the Morphology, Ontogeny and Embryology of the Flowers of Hedycarya Arborea J.R. et G. Forst. (Subfamily Monimioideae) and Laurelia Novae-Zelandiae A. Cunn. (Subfamily Atherospermoideae) of the Family Monimiaceae

Abstract

The inflorescences, flowers and the vascularization of floral parts of Hedycarya arborea and Laurelia novae-zelandiae were described and comparisons made with other members of the family in an attempt to determine the basic types of inflorescences, flowers and floral vascularization in the family. The vegetative, inflorescence and floral meristems of the two genera were compared. It was concluded that the vegetative apices of both had the tunica-corpus configuration typical of many other woody Ranales and other orders. The inflorescence apices were quite similar to the vegetative ones. The young floral apices are in a state of transition from a tunica-corpus to a mantle-core configuration and older floral apices had the mantle-core configuration, which is typical of the floral apices of many woody Ranales. Unusual features of the floral apices of Hedycarya and Laurelia were the lack of a pronounced rib meristem and the occurrence of relatively frequent divisions within vacuolate cells of the core. The ontogeny of the stamens of Hedycarya and Laurelia was described and comparisons were made. In both genera the micro-sporangium developed in a similar fashions: in Hedycarya 5-6 wall layers are formed inside the epidermis; in Laurelia there are 3-5 layers. Both genera had a typically thickened endothecium and a tapetum of the secretory type in which the tapetal cells become binucleate during the first meiotic division of the pollen mother cells. In Hedycarya the meiotic divisions of the pollen mother cells are of the successive type in which walls form by means of centrifugal cell plates Pollen grains remain in permanent tetrads in this genus. In Laurelia wall formation at the end of meiosis is of a modified simultaneous type, which may not have been hitherto described in the literature. Pollen grains are not in permanent tetrads. When the first division occurs in each microspore in Hedycarya, all four cells of a tetrad are at the same stage of division and the generative cell is cut off towards the distal face of the grain. Each microspore is in the two celled condition when shed. It was deduced that the generative cell is cut off against what represents a radial wall of the grain (with reference to the tetrad stage) in Laurelia. Pollen is shed in either the two or three celled condition. Comparisons were made with the development of microsporangia and male gametophytes in other woody Ranales. A study was made of the ontogeny, structure and function of the staminal appendages of Laurelia. It was found that the appendages function as nectaries, the nectar being predominantly sucrose. After a discussion of the various theories as to the morphological nature of the staminal appendages of the Laurales, it was concluded that they are morphologically staminodes. The carpels of Hedycarya and Laurelia have a basically similar ontogeny in which, as in the Lauraceae, the terminal stigmatic region develops from a solid terminal meristem in contrast to many woody Ranales in which the stigma-consists of crests which surround the external part of the cleft of the carpel. The ovules of Hedycarya and Laurelia resemble those of most other woody Ranales in being bitegmic, crassinucellate and anatropous with a monosporic 8-nucleate embryo sac of the Polygonum type. Both linear and T-shaped megaspore tetrads were found in the two genera. Laurelia has pseudocarps which develop after anthesis and enclose plumose achenes, but in Hedycarya the fruits are drupes. It was concluded that Laurelia and Hedycarya belong to two subfamilies which have been separated from each other for a long time and have undergone considerable evolution in different directions. It was also concluded that the Monimiaceae are closely related to the Lauraceae.