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The Geography of Phytochemical Races

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138 3 After the Ice<br />

No geographic structure was revealed by this analysis, with trees from Arch Cape,<br />

Oregon, Whitefi sh, Montana, and sites from northern British Columbia, including<br />

Queen Charlotte Islands, being closely associated. <strong>The</strong> authors remarked on the<br />

lack <strong>of</strong> differentiation between coastal and interior populations. A reinvestigation <strong>of</strong><br />

red cedar from 55 sites (3–6 trees per site) provided a new data set that was analyzed<br />

by numerical and discriminant-function analyses (von Rudl<strong>of</strong>f et al., 1988). <strong>The</strong>se<br />

analyses confi rmed the low intra- and interpopulational variation seen in the earlier<br />

study, but did reveal small differences between coastal and interior populations. No<br />

correlations between northern and southern populations emerged from the analyses;<br />

likewise, elevation had no effect on terpene composition.<br />

Parallel with the study just described was an examination <strong>of</strong> genetic variation in red<br />

cedar using enzyme electrophoresis (Yeh, 1988). Useful data from 2300 megagametophytic<br />

samples from 230 trees in eight populations were obtained for 15 enzymes representing<br />

19 loci. Sampling ranged between 48°49′ and 52°37′ N latitude and 116°40′<br />

and 124°10′ W longitude. <strong>The</strong> number <strong>of</strong> alleles per locus averaged 1.17 and observed<br />

heterozygosity ranged from 0.024 to 0.055 (average 0.036). <strong>The</strong> level <strong>of</strong> variability <strong>of</strong><br />

western red cedar approaches the lower limit for primarily outcrossing taxa (Hamrick<br />

et al., 1979) and is in accord with the very low levels <strong>of</strong> variation reported from the<br />

terpene analyses. Yeh (1988) suggested that the low levels <strong>of</strong> variation in red cedar, an<br />

outcrossing species that would be expected to be more variable, could be accounted<br />

for by comparatively rapid expansion following a bottleneck. He goes on to suggest<br />

that divergence time since the bottleneck is in the neighborhood <strong>of</strong> 10,000 years. It is<br />

known from the fossil record that red cedar did not reach the northern end <strong>of</strong> Vancouver<br />

Island until about 3000 years ago. Estimates <strong>of</strong> the timing <strong>of</strong> red cedar migration<br />

on the mainland range from 4900 to 10,000 years ago, which is consistent with the<br />

value obtained from the electrophoretic data. Further insights require information on<br />

effective population size and mutation rate per locus per generation (Yeh, 1988).<br />

3.1.6 Liriodendron tulipifera (Magnoliaceae)<br />

Liriodendron, tulip tree, consists <strong>of</strong> two species, the North American L. tulipifera L.,<br />

which we will examine here, and the Asian L. chinense (Hensley) Sarg. <strong>The</strong> relationship<br />

between the two species will be discussed later in this review. <strong>The</strong> North American<br />

tulip tree occurs naturally throughout a wide range in eastern North America from<br />

28°N to 43°N latitude, predominantly east <strong>of</strong> the Mississippi River. It is also commonly<br />

planted as a decorative species in many parts <strong>of</strong> North America (it fares well in<br />

Vancouver, B.C. at 49°13′N). In its natural setting, the tulip tree occurs in two forms,<br />

an upland form that can occur in large populations in the northerly part <strong>of</strong> its range,<br />

and an ecotype usually found in low, wet habitats along the coastal plains from North<br />

Carolina to Louisiana (Parks and Wendel, 1990, and citations therein; Parks et al.,<br />

1994). <strong>The</strong> latter report described an electrophoretic study <strong>of</strong> allozyme variation across<br />

the range <strong>of</strong> the species, from which three forms emerged based on differences in

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