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642 <br />0.55 <br />U 0.4 <br />a <br />COPEIA, 1995, NO. 3 <br />0 <br />0 0 <br />0 0 00 0 <br />° 0° ° <br />• • •• • 0 0 0 <br />•o ?• 'Op 8000 ° <br />M 0 00 o °to °do % o <br />Mb A A o °X 180 <br />omOYd am(s) <br />• 0 a °°°QL0 <br />% <br />• <br />r <br />0.25 4- <br />0.02 <br />0.15 <br />PC 11 <br />a <br />4 <br />2 <br />U 0 <br />-2 <br />0.28 -4 <br /> <br />?A <br />0 <br />v o?o? o a • <br />o • e• as :0 <br />0 A <br /> <br />? <br />O• 641 *A <br />• <br />, <br />° <br />?'j <br />3 <br />o? <br />? <br />®? •• • <br />• O <br /> <br />C6 0 <br />_ <br />A <br />o <br />V, o <br />y <br />•? <br />tt <br />o ?'?S ? y <br /> <br />o • % <br />-8 -6 -4 -2 0 <br />CV 1 <br />U <br />C, D <br />C, D <br />Specimen <br />Fig. 6. Canonical variates analysis of morpholog- <br />ical variation within and between Gila robusta and G. <br />b cypha. CV I and II represent the first two shape factors <br />extracted from this analysis. There is a clear sepa- <br />ration of the following: (1) a G. robusta group (closed <br />circles) composed of specimens from Black Rocks, <br />Westwater and Debeque canyons, Rifle, and the Yam- <br />pa River; (2) a G. cypha group (open circles) repre- <br />senting samples from Black Rocks, Westwater and <br />Grand canyons, and the Yampa River; and (3) a mixed <br />group of G. robusta (open triangles) and G. cypha (closed <br />triangles) from Cataract and Desolation canyons. <br />Fig. 5. Principal component (a) and canonical var- <br />iates (b) analyses of morphological divergence be- <br />tween specimens of Gila robusta and G. cypha. In (a), <br />axes represent the first two shape factors derived from <br />the analysis. PCA provides evidence of structure be- <br />tween G. robusta (closed circles) and G. cypha (open <br />circles), but within-species point clouds overlap con- <br />siderably. In (b), the sample of G. robusta (specimens <br />1-215) clearly differs from that of G. cypha (specimens <br />216-363). Samples of both species taken from Cata- <br />ract and Desolation canyons (indicated by labeled bars) <br />are, on average, more intermediate (scores closer to <br />zero) than are other samples. <br />ciated with relative size and positions of the <br />dorsal and anal fins (ODo-OA, OA-vIA, OA- <br />IA) were characterized by having the largest <br />univariate F values. All specimens were cor- <br />rectly classified with DFA. <br />Geographic patterns of variation.-In addition to <br />clear separation of species, analyses of variation <br />at the generic level indicated the presence of a <br />strong locality effect on relationships among <br />several populations. The separation of groups <br />(e.g., populations) through canonical variates <br />analysis was good; 55 of 56 univariate character <br />ANOVAs displayed a significant population ef- <br />fect (P < 0.0009), and multivariate tests were <br />highly significant (ex. Wilk's Lambda = 0.0002, <br />F672,3510.7 = 2.314, P < 0.0001). As before, char- <br />acters describing fin placement (OA-vIA, ODo- <br />OA, OA-IDo) showed the largest univariate F <br />values. Nine of 12 canonical roots contained a <br />significant component of among-group varia- <br />tion. CVI and II produced a clear separation of <br />three groups: (1) a G. cypha group consisting of <br />individuals from Black Rocks, Westwater and <br />Grand canyons, and the Yampa River; (2) a G. <br />robusta group consisting of fish from Black <br />Rocks, Westwater and Debeque canyons, Rifle, <br />and the Yampa River; and (3) a mixed group <br />of G. robusta and G. cypha individuals from Des- <br />olation and Cataract canyons (Fig. 6). Greater <br />than 92% (336 of 363) of all specimens were <br />correctly classified using DFA. In addition, de- <br />spite an overall classification error rate of 0.075 <br />(27 of 363 misclassified), the majority (89%) of <br />misclassifications occurred within rather than <br />among species (Table 2). Three specimens of <br />G. cypha were assigned incorrectly to G. robusta <br />populations; no individuals of G. robusta were <br />classified as G. cypha. <br />Cluster analysis of generalized canonical dis- <br />tances among populations produced two pri- <br />mary, and somewhat contradictory, results. First, <br />conspecific populations tended to cluster to- <br />gether to the exclusion of heterospecifics (Fig. <br />7); samples of G. robusta and G. cypha generally