Ecological Wood Anatomy of the Woody Southern Californian Flora

Abstract

Wood of 207 species, representing all 178 woody genera of the Munz flora of southern California, was studied by means of sections and macerations. Data were gathered on features relating to the conducting system: number of vessels per mm2, diameter of vessels, length of vessel elements, number of bars per perforation plate, presence of true tracheids, vasicentric tracheids, vascular tracheids, helical sculpture, and growth rings. The occurrence of these features is analysed both with respect to each other and to ecological groupings and habit groupings. Statistically significant data permit ecological groupings to demonstrate degree of xeromorphy in wood features. Xeromorphy is indicated by more numerous vessels per mm2, narrow vessels, shorter vessel elements, presence of vasicentric tracheids or vascular tracheids, presence of helical sculpture on vessel walls, and presence of well-marked growth rings (growth rings are common in moist habitats because in southern California these are also montane and therefore cold in winter). All of these appear to have developed in many phylads independently. Vessel element length appears to change less rapidly, at least in some phylads (those with true tracheids) than the other features. Presence of scalariform perforation plates and of true tracheids is interpreted as relictual; scalariform plates occur virtually only in mesic habitats and in a small number of species. True tracheids, although relictual in nature, have been preferentially preserved because of the value of their enormous safety. Groups without true tracheids have evolved vasicentric tracheids or vascular tracheids (the three types are mutually exclusive) to a high degree. By deducting the species with true and vascular tracheids, one finds that 100% of the alpine shrubs, 77% of the desert shrubs, and 75% of the chaparral shrubs which could possibly have evolved vasicentric tracheids actually have them. These are the three ecological groupings which have vasicentric tracheids not only in southern California, but other areas of the world as well. Tracheid presence (and to a lesser extent vasicentric tracheid presence) forestalls vessel grouping, but in tracheid-free groups vessel grouping is a highly adaptive strategy for xeromorphy. One can rank xeromorphic connotation of qualitative features on the basis of data herein: growth rings are the most common numerically, followed by helical sculpture, vasicentric tracheids, and vascular tracheids. Vasicentric tracheids, like true tracheids, tend to occur in evergreen shrubs whereas vascular tracheids tend to be related to drought-deciduous shrubs. Among quantifiable features, number of vessels per mm2 changes more rapidly than vessel diameter. Scalariform perforation plates, true tracheid presence, and long vessel elements are associated with each other statistically . By entering number of woody species for each genus in the flora and performing appropriate computations, a figure for each feature is projected on the basis of the 512 woody species of southern California. This pro-rated figure shows that phylads with any of the mechanisms cited as signifying xeromorphy speciate much more rapidly than do the phylads with mesomorphic wood features.