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The Geography of Phytochemical Races

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6.4 Juan Fernandez Islands 265<br />

(fl avonol) derivatives were not seen in any <strong>of</strong> the species <strong>of</strong> subgen. Phoenicoseris.<br />

Other differences are relatively minor, accountable by operation (or lack there<strong>of</strong>)<br />

<strong>of</strong> a single enzyme. For example, all four species in subgen. Dendroseris appear<br />

to be unable to make apigenin diglucosides, but at least some <strong>of</strong> the members <strong>of</strong><br />

the other two subgenera can. However, without knowing precisely what the structures<br />

<strong>of</strong> these compounds are, it is impossible to draw any conclusions. It is useful<br />

to note, however, that each <strong>of</strong> the Masafueran species exhibits a fl avonoid pr<strong>of</strong>i le<br />

closely resembling at least one <strong>of</strong> the species from the respective subgenera from<br />

Masatierra. At most, there would appear to be single enzyme differences involved.<br />

In an effort to locate a likely, or at least possible, mainland ancestral type, these<br />

workers determined the fl avonoid pr<strong>of</strong>i les <strong>of</strong> representatives <strong>of</strong> a number <strong>of</strong> candidate<br />

genera. <strong>The</strong>ir results pointed to Hieracium and Hypochaeris as possibilities,<br />

with their choice going to a species <strong>of</strong> the former from Chile. More recent work<br />

using macromolecular techniques suggested closer ties to Sonchus s.l., or possibly<br />

Embergeria from New Zealand (Bohm, 1998b, p. 252; parenthetical remark<br />

added by T. F. Stuessy, volume editor). It is prudent to point out that, even with<br />

detailed fl avonoid chemistry <strong>of</strong> possible ancestral taxa, comparatively few additional<br />

insights would be gained owing to the simple and homogeneous fl avonoid<br />

chemistry <strong>of</strong> these genera (Bohm and Stuessy, 2001).<br />

Insights into evolutionary relationships <strong>of</strong> Dendroseris have been gained through<br />

application <strong>of</strong> macromolecular tools. <strong>The</strong> morphological studies <strong>of</strong> Sanders et al.<br />

(1987) provided the background for all subsequent investigations, fi rst by uniting<br />

in one genus the 11 taxa that had, prior to then, been dispersed in four genera, and<br />

then by providing the fi rst explicit phylogeny <strong>of</strong> the genus. Electrophoretic patterns<br />

within the genus were studied by Crawford et al. (1987), who found comparatively<br />

low level <strong>of</strong> genetic variation compared to other insular endemics (De Joode and<br />

Wendel, 1992). Esselman et al. (2000) described a study <strong>of</strong> random amplifi ed polymorphic<br />

DNA (RAPD) markers, while a study by Crawford et al. (1992a) involved<br />

a restriction-site analysis <strong>of</strong> chloroplast and nuclear ribosomal DNA. Although four<br />

species were missing from that work, owing to limited quantities <strong>of</strong> plant material,<br />

results were encouraging in that the emerging phylogeny was very similar to that<br />

obtained from the other data sets. <strong>The</strong> sampling problem was partially overcome<br />

when Sang et al. (1994) employed the polymerase chain reaction (PCR) technique<br />

to amplify DNA (using internal transcribed spacer (ITS) region <strong>of</strong> nuclear ribosomal<br />

DNA) making it possible to include two additional species. Again, subgenera<br />

Dendroseris and Phoenicoseris emerged as strongly supported clades. <strong>The</strong> situation<br />

with regard to members <strong>of</strong> subgen. Rea is less straightforward.<br />

6.4.2 Robinsonia (Asteraceae)<br />

Robinsonia (Asteraceae: Senecioneae), the second largest endemic genus on the<br />

islands, consists <strong>of</strong> seven species assorted into two subgenera, subgen. Rhetinodendron,<br />

consisting solely <strong>of</strong> R. berteroi (Hemsl.) Sanders, Stuessy & Marticorena

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