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The Geography of Phytochemical Races

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3.2 North American Conifers 141<br />

from the Texas mainland and Gulf Coast Islands and seven populations <strong>of</strong> the closely<br />

related A. cumanensis Kunth. from near Vera Cruz, Mexico, where both species grow<br />

sympatrically (Fig. 3.6). In addition to the sesquiterpene fractions, these workers<br />

also examined the volatile terpenes <strong>of</strong> these plants. Numerical analysis <strong>of</strong> the combined<br />

chemical data revealed a closer relationship <strong>of</strong> the island populations with the<br />

population from Vera Cruz than with populations on the Texas mainland. <strong>The</strong> earlier<br />

suggestion that the island was colonized from achenes originating from the mainland<br />

has to be discarded in view <strong>of</strong> the strong support for the parental populations lying<br />

farther south, possibly in the vicinity <strong>of</strong> Vera Cruz as suggested by the newer chemical<br />

data. It is thought that the islands originated within the last 5000 years as a result<br />

<strong>of</strong> post-Pleistocene sea level changes. Colonization <strong>of</strong> a newly formed island by<br />

means <strong>of</strong> a single achene, or a few achenes from a localized source, would explain<br />

the observed morphological homogeneity. <strong>The</strong> existence <strong>of</strong> populations on the<br />

mainland that exhibit the dilactone chemistry characteristic <strong>of</strong> island plants can be<br />

rationalized simply by transport <strong>of</strong> propagules to the mainland by prevailing winds<br />

during the time <strong>of</strong> year when A. psilostachya fruits are available.<br />

3.2 North American Conifers<br />

<strong>The</strong> next series <strong>of</strong> examples involve conifers whose secondary chemistry has been <strong>of</strong><br />

major interest for many years. <strong>The</strong> resulting literature in this area, particularly with<br />

regard to the application <strong>of</strong> terpenoids to taxonomic and evolutionary problems, is a<br />

large and rich one. In the paragraphs that follow, only a representative sampling <strong>of</strong><br />

the available examples will be described. An important early work is Mirov’s (1967)<br />

book <strong>The</strong> Genus Pinus, in which he summarized much <strong>of</strong> the available literature<br />

on the application <strong>of</strong> turpentine composition to unraveling relationships among the<br />

pines. Von Rudl<strong>of</strong>f’s 1975 review is also a valuable source <strong>of</strong> information.<br />

<strong>The</strong> use <strong>of</strong> volatile chemicals as systematic markers has the obvious advantage <strong>of</strong><br />

lending itself to quantifi cation through GLC. In many, if not most, <strong>of</strong> the cases discussed<br />

below, qualitative differences in monoterpene pr<strong>of</strong>i les would not have been<br />

suffi cient to allow distinctions to be made between taxa, or even between individuals<br />

within a population. This is true because most conifers synthesize many <strong>of</strong> the<br />

same monoterpenes, although <strong>of</strong>ten in vastly different relative concentrations. It is<br />

these quantitative differences that have been constructively used in the following<br />

examples. Structures <strong>of</strong> the terpenes commonly studied are presented in Fig. 3.7.<br />

3.2.1 Abies (Pinaceae)<br />

Abies—the true fi rs—comprise a moderately large genus, 49 species (Mabberley,<br />

1997, p. 1) that include some <strong>of</strong> the most important commercial species among the<br />

conifers, particularly in North America. Some <strong>of</strong> the more important North American

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