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The Geography of Phytochemical Races

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3.2 North American Conifers 159<br />

Foliage <strong>of</strong> Douglas fi r had only trace amounts <strong>of</strong> (+)-camphene in trees from the<br />

coast but major concentrations in trees from the other three sites. In contrast, terpene<br />

mixtures from lodgepole pine leaf and bole tissues had nearly the same levels<br />

<strong>of</strong> the major compound, (−)-β-phellandrene, and minor components regardless <strong>of</strong><br />

origin. Patterns <strong>of</strong> variation in the other species were less extreme.<br />

3.2.6 Sequoia sempervirens (Cupressaceae)<br />

Sequoia sempervirens (D. Don) Endl., the coast redwood, is one <strong>of</strong> the most striking<br />

members <strong>of</strong> coastal forests <strong>of</strong> the western United States. <strong>The</strong> range <strong>of</strong> the redwoods<br />

extends from southwestern Oregon to the Santa Cruz region <strong>of</strong> central California.<br />

Although restricted to a strip <strong>of</strong> coastal North America at the present time, fossil<br />

evidence shows that Sequoia was well developed in the Upper Jurassic and that the<br />

genus enjoyed a wide distribution in appropriate habitats throughout the Northern<br />

Hemisphere. Despite the importance <strong>of</strong> this species, both from the commercial<br />

forestry standpoint as well as a species <strong>of</strong> major tourist attraction, comparatively little<br />

work had been done on establishing limits <strong>of</strong> variation with the apparent exception <strong>of</strong><br />

varieties based upon leaf and branch morphology (Dallimore and Jackson, 1966).<br />

In order to rectify this state <strong>of</strong> affairs, Hall and Langenheim (1987) undertook a<br />

study <strong>of</strong> monoterpenes <strong>of</strong> the leaves <strong>of</strong> redwood collected from sites representing<br />

essentially the entire range <strong>of</strong> the species. This was done in two ways, sampling <strong>of</strong> 13<br />

wild populations and sampling <strong>of</strong> trees grown from seed collected at eight locations<br />

throughout the species’ range and grown and maintained under cultivation for about<br />

20 years at the Russell Reservation, Contra Costa County, California. Fifteen compounds<br />

were identifi ed and quantifi ed as percentages <strong>of</strong> the total monoterpene fraction.<br />

α-Pinene was the major monoterpene followed, in descending order, by the common<br />

terpenes limonene, sabinene, β-phellandrene, γ-terpinene, and myrcene. <strong>The</strong> other<br />

identifi ed compounds, α-thujene, camphene, β-pinene, α-phellandrene, α- terpinene,<br />

the cis- and trans- isomers <strong>of</strong> ocimene, p-cymene, and terpinolene, were present at levels<br />

too low to be <strong>of</strong> use in numerical analysis. In general, results from analysis <strong>of</strong><br />

wild-collected material and cultivated material were in agreement with similar levels<br />

<strong>of</strong> interpopulation variation which points to the source <strong>of</strong> variation being genetic rather<br />

than environmental.<br />

Cluster analysis revealed the existence <strong>of</strong> three groups <strong>of</strong> populations with a strong<br />

break in the vicinity <strong>of</strong> San Francisco Bay (Fig. 3.10). Population group 3 occurred<br />

exclusively south <strong>of</strong> San Francisco Bay, whereas the other two types occur scattered<br />

throughout the northern part <strong>of</strong> the range <strong>of</strong> the redwoods. Although the three population<br />

groups exhibited all six marker terpenes, obvious distinctions can be seen, even<br />

with visual examination <strong>of</strong> the data. Thus, all northern populations, that is, population<br />

groups 1 and 2, exhibited higher levels <strong>of</strong> α-pinene and β-phellandrene. <strong>The</strong><br />

highest levels <strong>of</strong> limonene, sabinene, myrcene, and γ-terpinene were observed in type<br />

1 populations, although the level <strong>of</strong> variation among populations was substantial.

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