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The Geography of Phytochemical Races

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202 4 Intercontinental Disjunctions<br />

ether [355] and 7,8,4′-trimethyl ether [356]. Eucryphia moorei exhibited a pr<strong>of</strong>i le<br />

somewhat simpler than that <strong>of</strong> E. jinksii but, as mentioned, lacked fl avones with the<br />

isoscutellarein oxygenation pattern. Eucryphia wilkiei exhibited a pr<strong>of</strong>i le consisting<br />

<strong>of</strong> a single, unidentifi ed aglycone, seen as well in E. lucida (Tasmania) and E. jinksii<br />

(mainland Australia). It was also unique in the genus in having fl avonol glycosides<br />

as exudate components.<br />

Numerical treatments <strong>of</strong> the fl avonoid data were performed using nonmetric<br />

multidimensional scaling (NMDS), and unweighted pair group method with arithmetic<br />

mean (UPGMA) or average linkage clustering <strong>of</strong> Bray-Curtis dissimilarities.<br />

<strong>The</strong> NMDS treatment resulted in clear separation <strong>of</strong> E. wilkiei, but the other six taxa<br />

appeared scattered, although geographically close pairs, for the most part, appeared<br />

closer to each other than to other species. <strong>The</strong> UPGMA treatment <strong>of</strong> Bray-Curtis<br />

values resulted in clear separation <strong>of</strong> E. wilkiei from all other species and the following<br />

pairings, E. cordifolia-E. glutinosa (Chilean species); E. jinksii-E. moorei<br />

(mainland Australia); and E. lucida-E. millinganii (Tasmania). Cladistic analysis<br />

<strong>of</strong> the fl avonoid data failed to resolve any relationships. A recent cladistic analysis<br />

using morphological data (Taylor and Hill, 1996) suggested that E. lucida and<br />

E. milliganii are sister taxa, which is supported by the phytochemical data. Neither<br />

the relationship <strong>of</strong> E. wilkiei to E. lucida and E. milliganii, as suggested by the cladistic<br />

analysis <strong>of</strong> Taylor and Hill (1996), nor to E. jinksii, as viewed by Forster and Hyland<br />

(1997), is supported by the fl avonoid data. Because <strong>of</strong> incongruities between relationships<br />

suggested by the phytochemical data and those that arose from cladistic<br />

analysis <strong>of</strong> morphological data, one is once again inclined to suggest that genesequence<br />

studies might be <strong>of</strong> value in this system. In addition to the obvious phylogenetic<br />

insights that might be gained from such a study, some ideas <strong>of</strong> what gains<br />

and losses have occurred in the fl avonoid biosynthetic pathway that resulted in the<br />

signifi cantly different pr<strong>of</strong>i les <strong>of</strong> compounds observed within the genus.<br />

4.4.2 Blennospermatinae (Asteraceae)<br />

<strong>The</strong> pattern <strong>of</strong> distribution <strong>of</strong> the four genera that comprise this subtribe is one <strong>of</strong><br />

the more complex ones that we encounter in this review, in that it involves western<br />

North America, South America, including the Falklands and the Juan Fernandez<br />

Islands; Australia, including Tasmania; New Zealand, including Stewart Island, the<br />

sub-Antarctic Campbell and Auckland Islands, and New Guinea. <strong>The</strong> four genera<br />

involved are the monotypic western North American Crocidium, Blennosperma with<br />

two Californian species, and one species disjunct in Chile, Ischnea with four species<br />

endemic to New Guinea (Swenson, 1994), and Abrotanella, which consists <strong>of</strong> 19 species<br />

distributed in the other areas listed. <strong>The</strong> subtribe has been the subject <strong>of</strong> recent<br />

biosystematic and geographic study (Swenson, 1995a, b; Swenson and Bremer,<br />

1997). From these data, it is clear that the subtribe represents a monophyletic group.<br />

In a paper describing pollen-wall ultrastructure, Skvarla and Turner (1966)<br />

stated that the fl avonoids <strong>of</strong> Blennosperma and Crocidium exhibited different

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