COMPARATIVE MORPHOLOGY AND PHYLOGENY OF THE RANALES

Abstract

SUMMARY Comparative morphological studies of woody Ranales have established the primitive status of the group and hence their key place in angiosperm phylogeny. Significant advances in our knowledge of some ranalian families have been made in recent years. An attempt is made in the present review to bring together a range of morphological data (vegetative and floral anatomy, palynology and embryology) on the Ranales (sensu lato), with particular reference to research work published after the publication of Eames's (1961) book, and to discuss the relationships of the families. Recent ontogenetic studies have shown that the carpel of Drimys is ascidial and not conduplicate as earlier suggested. The inclusion of Degeneria in the Winteraceae is not supported by morphological data. Melville's gonophyll theory has been shown to be inapplicable to the magnoliaceous flower. The pollen of Schisandra is interpreted as derived and specialized rather than primitive as previously supposed. The removal of Schisandra from Magnoliaceae is upheld by morphological evidence. Recent morphological studies do not support a close relationship between Schisandraceae and Illiciaceae suggested by earlier authors. The Canellaceae shows similarities to Winteraceae, Magnoliaceae, Illiciaceae, Eupteleaceae and Myristicaceae. Transitional types of division of pollen mother cells found in Winteraceae, Schisandraceae and Annonaceae and their probable phylogenetic significance have been discussed. The Annonaceae, Winteraceae, Degeneriaceae, Magnoliaceae, Schisandraceae and Cercidiphyllaceae share several embryological features in addition to similarities in floral structure. Ruminate endosperm is regarded either as an archaic feature retained in some taxa or as a later and parallel development in others. Thus its value in assessing relationships seems to be doubtful. Myristicaceae has been shown to be closely related neither to the the Annonaceae nor to the Lauraceae. The suggested relationship of Eupomatiaceae to Annonaceae is not supported by palynology. Floral cortical vascular systems in Magnoliaceae, Annonaceae, Calycanthaceae and Myristicaceae have been compared and it is concluded that they may be vestigial structures. A great deal of similarity has been found between Lauraceae and Calycanthaceae in wood, node, flower structure and embryology. Further floral anatomical evidence has been adduced to support the removal of Scyphostegia from Monimiaceae. The Hernandiaceae show similarities to some members of Monimiaceae while the Gyrocarpaceae resemble the Lauraceae, Gomortegaceae and certain other genera of Monimiaceae. Available evidence from wood and floral anatomy and embryology indicates close relationships among Lauraceae, Monimiaceae and Hernandiaceae. Vegetative and floral anatomical and embryological data seem to indicate a place for the Chloranthaceae in the ranalian complex. Recent anatomical studies in the Nymphaeaceae show that the floral structure is of a primitive type with similarities to the woody Ranales. Available morphological evidence is considered inadequate to express an opinion on the splitting of the family. Ceratophyllaceae is regarded as a highly reduced ranalian family derived most probably from a nymphaeaceous stock. The gynoecium in Berberidaceae is interpreted as monocarpellate. No evidence has been found to support the tricarpellate view. Berberidaceae, Lardizabalaceae and Menispermaceae share several embryological features, while at the same time showing evidence of specialization, each in its own way. Thus they might have arisen from a common stock and early diverged along different lines. The occurrence of several types of embryo sac in Ranunculaceae may well be an indication of specialization, but their probable taxonomic value, if any, is not yet clear. The occurrence of numerous primitive features in Paeonia has been suggested as an argument for its retention in the Ranales. No evidence has been found to preclude the inclusion of Dilleniaceae in the Ranales. On the other hand, as opposed to similarities in wood and pollen characters between Dilleniaceae and Theaceae, floral anatomical and embryological features offer a sharp contrast between the two. The Ranales are believed to be polyphyletic. It has been tentatively suggested that two major phyletic lines may be recognized in each of the woody and herbaceous series: the magnolialian and lauralian lines in the former and the nymphaealian and berberidalian lines in the latter.

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