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McELROY AND DOUGLAS-MORPHOLOGICAL VARIATION IN GILA 643 <br /> <br /> <br />were found to be distinct morphologically both <br />in sympatry and in allopatry. Second, at both <br />Desolation and Cataract canyons, heterospecific <br />populations were more similar to one another <br />than they were to conspecific populations from <br />other localities (Fig. 6, 7). Populations at these <br />sites tended to group by locality rather than by <br />taxonomic affinity. Results of UPGMA cluster- <br />ing suggest that this morphological conver- <br />gence is a within-locality (as opposed to a his- <br />torical) phenomenon; the node separating the <br />Desolation and Cataract canyon clusters is deep <br />(Fig. 7). <br />DISCUSSION <br />Data presented here demonstrate significant <br />among-group structure in morphological vari- <br />ation for G. robusta and G. cypha at three levels <br />of biological organization. Intraspecific analy- <br />ses indicate that population divergence exists in <br />both G. robusta and G. cypha; however, geo- <br />graphic proximity alone is insufficient to ac- <br />count for the hierarchical relationships ob- <br />served among populations of either species. <br />Comparison between species suggests that G. <br />robusta and G. cypha represent distinct mor- <br />phologies, both in sympatry and in allopatry. <br />Nevertheless, patterns of variation at the ge- <br />neric level indicate that heterospecific popula- <br />tions at Desolation and Cataract canyons are <br />morphologically more similar to one another <br />than to allopatric conspecifics. These results <br />have implications for the identification of evo- <br />lutionary and conservation units, the potential <br />Black Rxks <br />Yampa River <br />WesW_' Cr. anG-b- <br />Debegw Canyon <br />Rifle <br />Desolation canyon <br />Desolation Canyon <br />Camel canyon <br />Caramel canyon <br />Black Rocks <br />Wens -Canyon <br />G. ryyM <br />c.ana canymr <br />Yampa Riva <br />56.0 48A 40.0 32.0 24.0 16.0 8.0 <br />Fig. 7. Hierarchical relationships among sympat- <br />ric and allopatric populations of Gila robusta and G. <br />cypha. Dendrogram represents UPGMA clustering of <br />generalized-squared Mahalanobis distances among <br />group means. A strict consensus topology based on <br />UPGMA, complete linkage, and single linkage clus- <br />tering is nearly identical; a single branch (indicated <br />by a dashed line) collapses in the consensus topology. <br />effects of hybridization and local adaptation on <br />morphology, and the design of conservation <br />strategies for these fishes. <br />Population divergence.-Previous studies of mor- <br />phological variation among Gila from the upper <br />Colorado Basin (Smith et al., 1979; Douglas et <br />al., 1989; Kaeding et al., 1990) have been con- <br />cerned with partitioning morphological varia- <br />tion at the species level. In general, these studies <br />TABLE 2. CLASSIFICATION OF Gila robusta AND G. cypha SPECIMENS BY SPECIES AND POPULATION. The overall <br />classification error rate was 0.075; however, 89% of misclassifications occurred within (error rate = 0.066) <br />as opposed to among (error rate = 0.008) species. Rows represent actual group membership; population labels <br />are the same as for columns. Labels for each population are as in Table 1. <br /> <br /> <br />B <br />G. robusta <br />C D Q R <br /> <br />w Predicted group <br /> <br />y B <br />G. cypha <br />C D G w y <br />B 18 - - - 1 - - - - - - - - <br />C - 6 - - - - - - - - - - - <br />D - - 24 - - - - - - - - - - <br />Q - - - 19 1 - - - - - - - - <br />R - - - 1 24 - - - - - - - - <br />W 3 - - 1 - 51 1 - - - - - - <br />Y 1 - - 3 - 2 59 - - - - - - <br />B - - - 1 - - - 21 - - - 3 - <br />C - - - - - - - - 11 - - - - <br />D - - 1 - - - - - - 22 - - - <br />G - - - - - - - - - - 26 2 - <br />W - - - - - 1 - 5 - - - 51 - <br />I - - - - - - - - - - - - 5