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The Geography of Phytochemical Races

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116 2 Examples Within Continents<br />

Chrysosplenium fi ts into several phytogeographic categories. In addition to the<br />

local infraspecifi c differentiation just described, it also falls into the category <strong>of</strong><br />

genera with an eastern Asia–eastern North America–western North America distribution.<br />

For example, several members <strong>of</strong> the opposite-leafed species <strong>of</strong> Chrysosplenium<br />

occur in Japan with the group represented in North America by two<br />

species, C. americanum Schwein. ex Hook., which occurs along the eastern seaboard<br />

<strong>of</strong> North America, and C. glechomaefolium the western North American representative.<br />

Evolutionary relationships between these two North American species<br />

were discussed by Soltis et al. (1993) as part <strong>of</strong> a study <strong>of</strong> the genus using DNA<br />

sequence data. Included in that study were two species native to extreme southern<br />

South America, which will be discussed below under the category <strong>of</strong> polar disjunctions.<br />

Eastern Asian–eastern North American disjunct taxa will also be discussed<br />

later in this review.<br />

2.7.20 Quercus rubra (Fagaceae)<br />

Quercus rubra L., red oak, is a major component <strong>of</strong> the deciduous forests <strong>of</strong> eastern<br />

North America, with a range that extends from Ontario south to the Gulf States<br />

(Sargent, 1965). Part <strong>of</strong> the variability <strong>of</strong> the species has been formalized by the<br />

recognition <strong>of</strong> var. borealis (Michaux f.) Farwell to account for trees in the northern<br />

part <strong>of</strong> the species’ range, and at higher elevations in the southern Appalachians.<br />

<strong>The</strong> two varieties, rubra and borealis, are thought to intergrade between 1200 m and<br />

1350 m in the southern Appalachians.<br />

An examination <strong>of</strong> the fl avonoids <strong>of</strong> red oak by McDougal and Parks (1984)<br />

revealed altitudinal differences in the fl avonol pr<strong>of</strong>i les. One chemotype, characterized<br />

by the presence <strong>of</strong> kaempferol [234] and quercetin 3-O-glycosides [235],<br />

occurred in trees from lower elevations, whereas in trees from higher elevations<br />

myricetin 3-O-glycosides [236] (see Fig. 2.71 for structures 234–236) were added to<br />

the array. Mixed populations were observed between 600 m and 1050 m, somewhat<br />

lower than the elevational range <strong>of</strong> overlap <strong>of</strong> the morphological varieties known to<br />

grow between 1200 m and 1350 m (Sargent, 1965). <strong>The</strong> infl uence <strong>of</strong> environmental<br />

and genetic factors on the expression <strong>of</strong> fl avonols in red oak was the subject <strong>of</strong> a<br />

second paper by McDougal and Parks (1986). Acorns were collected from eight<br />

sites representing an elevational gradient <strong>of</strong> 1455 m, divided equally, and planted in<br />

nursery plots at 75–1140 m elevation. After 5 years, leaf samples were analyzed for<br />

fl avonoids qualitatively and quantitatively. Regardless <strong>of</strong> the elevation <strong>of</strong> the nursery,<br />

seedlings originating from high-elevation myricetin-positive trees retained their<br />

capacity to synthesize myricetin, while seedlings from low-elevation nonmyricetin<br />

stock continued to produce only kaempferol and quercetin derivatives. Quantitative<br />

analyses indicated that the total amounts <strong>of</strong> fl avonoids varied somewhat with elevation,<br />

but the array <strong>of</strong> compounds produced was not affected. Although the fl avonoid<br />

chemotypes correspond to the two varieties <strong>of</strong> red oak in the southern Appalachians,<br />

the zone <strong>of</strong> overlap <strong>of</strong> chemical and morphological characters does not agree.

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