Relationships of the Centrospermae

In the past, most systematists accepted the Polygona-ceae (occasionally also the Plumbaginaceae) as the closest relatives of the Centrospermae. This idea had already been suggested by R. Brown (1810), based on similarities such as the uniseriate perianth, the unilocular ovary, the single ovule, and the basal placenta. However, the unilocular and uniovulate ovary has probably evolved within the Centrospermae and is therefore no proof of a common descent with the Polygonales. On the other hand, some striking similarities exist between the Centrospermae and the Polygonales: similar sequences of cytochrome C were found by Boulter et al. (1972) and of other macro-molecules by Martin et al. (1985); and certain rust fungi occur on plants of both orders and of the Plumbaginaceae (Savile 1979). However, the hydrolyzable tannins occurring in Polygonaceae and Plumbaginaceae remove them strongly from the Centrospermae.

After recognizing the phenomenon of centrifugal stamen development, Cronquist (1965) and similarly Thome (1976) used it as a criterion for indicating a relationship between the dilleniid "Guttiferales" and the Centrospermae. While Cronquist (e.g., 1981) has long abandoned this view, Leins (1975) and Erbar (1986) still adhere to it, interpreting fasciculate an-droecia as plesiomorphic. However, since most Dille-niidae, in contrast to the Centrospermae, are basically woody, the search for a sister-group of the Centrospermae should focus on a predominantly herbaceous group. The rich representation of galloyl esters in the Dilleniidae, which have never been found in the Centrospermae, makes a relationship between them doubtful.

The Centrospermae show remarkable resemblances in various features to the monocotyledons. These include the presence of bound ferulic acid in unlignified cell walls; a dermal origin of the outer integument (present also in Theales: Bouman 1984); and the biogenetic relationship of betalains to alkaloids in the monocotyledons (see p. 23). Kendrick and Hillman (1971) found that seedlings of species of the Amaranthaceae, Chenopodiaceae, and Nyctagina-ceae, like members of the monocotyledons, lack the phytochrome dark reversion. It is likely that both the Centrospermae and the monocotyledons are basically herbaceous and returned only secondarily to woody growth (e.g., Cronquist 1981,1988). However, the phylogenetic tree constructed by Wolfe et al. (1989) based on the comparison of cpDNA sequences of four monocot and three dicot taxa makes a close relationship of the Centrospermae and monocotyledons very improbable.

A ranalean affinity of the Centrospermae is favored by many authors, including Takhtajan (1959), Roland (1969), van Campo (1976), Nowicke and Skvarla (1979), and Cronquist (1981). This suggestion is based mainly on the correspondence of pollen characters and the predominance of herbaceous growth in both groups. Takhtajan (1959), focusing on their anomalous secondary growth, the curved embryo, and gynoecium morphology, singled out the Menispermaceae as close relatives, Buxbaum (1961) favored the Illiciaceae because of the now discarded similarity of the gynoecia. The Illiciales and Berberi-

dales were considered by Huber (1982, 1990) as closely related to the Centrospermae, the former because of their unilacunar nodes, and the latter because of their mostly herbaceous life forms, anomalous growth, nuclear endosperm development, and the occasional presence of triterpene saponins. Further indicators of a ranalean affinity might be sought in the similarities in seed coat anatomy between the Centrospermae, Ranunculaceae, and Papaveraceae (Netolitzky 1926) and the ample utilization of tyrosine in secondary metabolism. The latter feature extends into the monocotyledons as well. On the other hand, while sieve-tube plastids with proteinaceous inclusions occur in many lower Magnoliidae and all monocotyledons, the Ranunculales have only starch in their plastids.

It is evident from this survey that only a broad analysis based on a large number of characters and making use of the parsimony principle will provide substantial progress in the problem of relationships of the Centrospermae.

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