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The Geography of Phytochemical Races

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88 2 Examples Within Continents<br />

<strong>The</strong> overall mean number <strong>of</strong> steps per population is 19.53 and the overall mean<br />

MBSD is 14.12. Comparative mean values for the separate ranges <strong>of</strong> P. carolina are:<br />

continuous range = 13.14 ± 4.95 (N = 2016), disjunct range = 15.42 ± 6.40 (N = 36),<br />

and continuous versus disjunct populations = 17.47 ± 3.90 (N = 576). <strong>The</strong>se differences<br />

are signifi cant (P < 0.001). <strong>The</strong>se values indicate that the separate ranges <strong>of</strong><br />

the species are distinctive, but only weakly divergent with regard to their fl avone<br />

pr<strong>of</strong>i les. Although there are no discrete patterns <strong>of</strong> fl avonoid pr<strong>of</strong>i les, some areas in<br />

the continuous range show higher pr<strong>of</strong>i le uniformity than others. Of particular note<br />

are populations in western Georgia, where similarities in the range 85–100% were<br />

observed. However, even in that area, adjacent populations may not be the most<br />

closely related, and those that are closely related may be separated by some distance<br />

(ca. 50 miles). Although differences in pigment pr<strong>of</strong>i les were found within this species,<br />

no clear-cut pigment races exist.<br />

This fi nding is in general accord with aspects <strong>of</strong> the biology <strong>of</strong> the species. Levy<br />

and Fujii (1978) pointed out that restricted gene fl ow is characteristic <strong>of</strong> Phlox,<br />

which lacks the capacity for long-distance dispersal. <strong>The</strong>y pointed out that the irregular<br />

occurrence <strong>of</strong> most <strong>of</strong> the compounds suggested that the gradients observed<br />

represent the results <strong>of</strong> “range dissection and contraction rather than stepping-stone<br />

dispersal from central or local centers <strong>of</strong> fl avone diversity.” <strong>The</strong> result would be a<br />

collection <strong>of</strong> local “islands” <strong>of</strong> fl avone diversity, which, through local expansion,<br />

might come into contact with nearby populations to produce the complex array <strong>of</strong><br />

variants seen.<br />

This study <strong>of</strong> Phlox carolina represents one <strong>of</strong> the best examples in the fl avonoid<br />

chemosystematic literature where workers combined thorough sampling, detailed<br />

statistical analysis, and an intimate knowledge <strong>of</strong> the biology <strong>of</strong> the system under<br />

scrutiny to produce a convincing picture <strong>of</strong> natural variation. As mentioned at the<br />

beginning <strong>of</strong> this discussion, a more casual approach would have undoubtedly overlooked<br />

the subtle differences that characterize this system.<br />

2.7.5 Lasthenia (Asteraceae)<br />

Lasthenia Cass. is mostly a Californian genus <strong>of</strong> about 17 species; the single species<br />

that occurs in Chile will be dealt with in the section on north–south disjuncts<br />

below. An examination <strong>of</strong> fl avonoid pigments <strong>of</strong> all members <strong>of</strong> the genus (Bohm<br />

et al., 1974) had shown that sections, defi ned on the basis <strong>of</strong> morphology, cytology,<br />

and breeding behavior (Ornduff, 1966), could also be distinguished on the basis <strong>of</strong><br />

the presence or absence <strong>of</strong> a variety <strong>of</strong> fl avonoids (Bohm et al., 1974). For example,<br />

sections Baeria (both species), Burrielia (all three species), and Hologymne (two<br />

<strong>of</strong> three species) were characterized by an array <strong>of</strong> chalcone and aurone derivatives<br />

(collectively referred to as anthochlors), in addition to a number <strong>of</strong> common<br />

fl avones and fl avonols, whereas members <strong>of</strong> the other three sections exhibited only<br />

fl avonols. <strong>The</strong> initial studies suggested that each section was characterized by a<br />

unique combination <strong>of</strong> pigments. Some indication <strong>of</strong> variation did emerge from

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