Taxus sumatrana Group

©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
April 2003, Feb. 2007

Mid leaf sections representative of the Taxus Sumatrana Group, drawn by R. Spjut.  From top to bottom: Upper two are T-sections of T. mairei, top: from Guangdong (China), Tsang 20694 (US), showing elevated truncate midrib on abaxial surface that has slightly larger epidermal cells than those on adaxial surface, the lower section from Yunnan, type (A), showing channeled midrib and larger spherical parenchyma cells lying against smaller epidermal cells— most conspicuous along midrib and marginal zones.  Note elliptically shaped epidermal cells.  Lower two sections: abaxial epidermis from margin to across midrib; upper most drawn from type of T. speciosa, showing marginal region of 23 smooth cells in width, a stomata band with 16 rows of stomata, and a smooth midrib, the lowest section from type of T. sumatrana, showing long rectangular cells and 12–14 stomata rows

 

The Sumatrana Group is characterized by leaves having a relatively broad region of marginal cells that border stomata bands, usually from 8–36 cells across. This is often evident in dried specimens by a glossy reddish discoloration.  The epidermal (accessory) cells in stomata bands are further differentiated from marginal and midrib regions by their shorter length and broader width.  Leaves in the other species groups usually have stomata bands bordered by fewer cells, or are less differentiated by color and development of papillae.

The taxonomy of this Sumatrana Group is difficult due to taxa having partially correlated character traits.   Previous taxonomists have recognized only one taxon in this group, either as a variety (Cheng & Fu 1975, 1978; Li & Fu 1997), or species (Florin 1948; Handel Manzzetti 1929; Hu 1964), but they have not applied the correct name.  For example, de Laubenfels (1978) applied the name T. sumatrana to all yews in SW Asia, but this antedated by T. wallichiana.  Further, this very broad concept of T. wallichiana makes it difficult to distinguish T. cuspidata and its allies (Baccata group).

For practical reasons I recognize more than one species in the Sumatrana Group based on differences in the leaves. The abaxial leaf surfaces of T. sumatrana and T. mairei usually have truncated midribs that are slightly channeled in contrast to rounded (keeled) midribs in T. kingstonii.  Taxus mairei is then differentiated by the abaxial midrib having short trapezoidal, inflated epidermal cells, compared to nearly rectangular cells in T. sumatrana (L-sect.), but young leaves of T. mairei may have nearly rectangular cells; thus, I regard the inflated short epidermal cells as the derived (apomorphic) trait.  Taxus celebica is primarily identified by the lanceolate acuminate shape of leaves that usually remain green and flat upon drying, whereas T. sumatrana has darker colored leaves—green to blackish green—that become puckered and revolute along margins upon drying.  Seed shape and seed color, which are variable in this group, could prove valuable to species delimitation; unfortunately, most specimens lack seed. Taxus speciosa Florin is treated as a variety of T. mairei because of their close similarity in leaf anatomy and geographical distribution.

Leaf anatomical differences in other species groups seem less significant because possibly of more frequent hybrids between temperate and subtropical species, although geographical differentiation of taxa is evident in other morphological characters.  For example, zigzag isodichotmous branching characterizes T. mairei in SE China (Fig. 63) in contrast to relatively short unequally much-divided branchlets of T. biternata in NE China and adjacent temperate areas (Figs. 145, 268).   This is in further contrast to longer divaricate branchlets of T. umbraculifera var. microcarpa in SE Russia (Fig. 165).  These differences appear to be a product of adaptive radiation with branchlets becoming more unequally divided from west to east.  Similarly, T. sumatrana in Indonesia and T. biternata (Baccata Group) in Japan may be synapomorphic with regard to their two ranked leaf arrangement and elliptical shape of epidermal cells.  The most logical time for diversification in branching and leaf arrangement is during the Paleocene when the northern latitudes possibly supported a subtropical flora (Hsü 1983; Manchester 1999; Wolf 1975).  On the other hand, a closer similarity in branching between some specimens of T. umbraculifera var. microcarpa in SE Russia (Fig. 288) and T. mairei in its northeastern most range may relate to hybridization or a later history of geographic and climatic differences since the Miocene.