Anatomy of the Secondary Phloem in Winteraceae

The secondary phloem of nine species in five genera of Winteraceae was examined with regard to features that could serve for taxonomic and phylogenetic evaluation of the family. The species examined were as follows: Bubbia pauciflora, B. semecarpoides, Drimys lanceolata, D. winteri, Exospermum stipitatum, Pseudo wintera axillaris, Zygogynum baillonii, Z. bicolor, and Z. vinkii. The nine species showed the following common characteristics: 1) origin from nonstoried vascular cambium with long fusiform initials; 2) ray system consisting of high multiseriate and high uniseriate rays; 3) occurrence of secondary partitioning in the differentiating phloem so that the sieve elements are much shorter than the tracheids; 4) lack of sharp differentiation between lateral sieve areas and those of the sieve plates; 5) predominance of compound sieve plates; 6) short companion cells, often single in a given sieve element; 7) phloem parenchyma cells in strands; 8) lack of specialised fibres (bast fibres) in the secondary phloem; 9) presence of nondispersing protein body in the sieve element protoplast. Features numbered 1, 2, 4-6 are considered to be indications of low evolutionary level. The significance of the other three features (3, 7-9) requires further evaluation. Among these three is the secondary partitioning the occurrence of which seems to imply that in some taxa the well known sequence of evolutionary shortening of cambial initials and their derivatives may be accelerated on the phloem side.