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<br />The distribution of haplotypes strongly reinforces the pattern detected by the F statistics. The <br />San Juan system is unique and the other subbasins are more similar to their adjacent basins (the <br />Colorado River subbasin to the Dolores River subbasin and the White River subbasin to the <br />Yampa River subbasin), yet they are still unique from one another in the ratios of the dominant <br />haplotypes. As has been noted earlier, the unique minor haplotypes tend to be most abundant in <br />the individual populations. The isolating mechanism generating this pattern appears to be both <br />distance and time. The presence of unique minor haplotypes within populations supports time as <br />an important factor because these unique forms have not spread to other parts of the basin. <br />Unfortunately we did not survey the speckled dace haplotypes in the subbasins to the Colorado <br />and Green Rivers within Utah, but it is likely that haplotype frequencies in other portions of the <br />drainage basin will further reinforce the observed patterns. The Gunnison system would be <br />expected to be much closer to the Colorado River subbasin than is the Dolores subbasin. <br />Unfortunately low sample size and lack of additional sampling sites does not allow us to address <br />this relationship. <br /> <br />Sequence similarity was also examined using the Tamura-Nei index (Tamura and Nei 1993). <br />MEGA (Kumar et a12001) was used to generate clusters of the similarity matrix (Figures 3a, <br />3b). Because this is a similarity based index, populations and subbasins can be joined in a <br />synthetic index, regardless of individual evolutionary trajectories. Thus multiple unique <br />haplotype sequences within a population are combined with haplotypes shared with other <br />populations to generate the synthetic stand for a given location. This allows a visual <br />representation of the overall similarity of populations and subbasins. The relationships between <br />basins are apparent in Figure 3a. The San Juan is the most dissimilar of the subbasins examined <br />in the upper Colorado Basin. The remaining subbasins are more similar to one another, with the <br />Colorado River subbasin being the most distinct, followed by the Gunnison and Dolores. The <br />White and Yampa are the most similar subbasins. <br /> <br />The associations among drainages are more clearly shown in the cluster of relationships among <br />sites (Figure 3b). The San Juan populations are most distant from other speckled dace <br />populations, and interestingly, the San Juan populations tend to show more similarity to the <br />overall cluster than to each other. They do not form a separate cluster of their own. The <br />Colorado, Gunnison and Dolores populations are quite similar to one another. Slight changes in <br />frequencies of various haplotypes would easily shift relationships among these populations. <br />Both the White and the Yampa populations form clusters where populations within the subbasins <br />are more similar to one another than to the adjacent subbasin. One exception to this is the <br />Piceance Creek sample from the White River (WhPi; Figure 3b; site #9), which is clustering <br />basally with the Yampa subbasin populations. This population (site # 9 on Figure 2), was pulled <br />to the Yampa subbasin (Figure 3) because of the high proportion of haplotype K (haplotype # 54; <br />Table 9). The population distance cluster does show the White and Yampa river populations as <br />being most similar, but they clearly have separate evolutionary trajectories. <br /> <br />Phylogenetic relationships among the speckled dace populations were examined using PAUP <br />(Phylogenetic Analysis Using Parsimony; Swofford 1998). We used longnose dace as our out <br />group, and included speckled dace from the Virgin River and Gandy marsh (Utah) to help anchor <br />the relationships. We identified 96 speckled dace haplotypes within the Colorado River <br /> <br />36 <br />