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1730 D. B. OSMUNDSON ET AL. Ecological Applications <br />Vol. 12, No. 6 <br />3.5 <br />3.0 <br />2.5 <br />c 2.0 <br />O <br />U 1.5 <br />S 1.0 <br />0.5 <br />0 <br />• Run AA a <br />o Riffle • tfo 0 <br />• °° <br />00•tt ®8 <br />• • 0 <br />• o• 00 <br />•• 0 0 <br />• <br />• • • 00 <br />• <br />• <br />-6 <br />4.5 <br />N 4.0 <br />3.5 <br />L? 3.0 <br />m 2.5 <br />9 2.0 <br />m <br />1.5 <br />c 1.0 <br />0.5 <br />0 <br />-5 -4 -3 -2 -1 0 <br />Principal component score <br />1 2 <br />° b <br />0 <br />°0 <br />° o <br />•o <br />• <br />• • •g o <br />0a® 0 <br />019 <br />• 0 oie <br />• <br />••w ° •0 <br />• <br />3.5 <br />a 3.0 <br />c 2.5 <br />2.0 <br />U <br />1.5- <br />1.0- <br />0.5 <br />0 <br />0 <br />3.5 <br />co <br />0 3.0 <br />a) 2.5 <br />aa) 2.0 <br />V_ <br />a> 1.5 <br />C <br />1.0 <br />U <br />0.5 <br />0 <br />U.5 I.U - 1.3 Z.U Z.D 3.U 15.3 <br />In(Measured chlorophyll a) <br />d <br />• <br />0 0 <br />• •• o <br />.0 • <br />• 0 0°O 00 0?4 <br />• •• • ° ° <br />• <br />• <br />-6 -5 -4 -3 -2 -1 0 1 2 0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 <br />Principal component score In(Measured invertebrates) <br />FIG. 4. Interrelationships among physical and biological parameters. Solid lines represent regression lines in (a) and (b) <br />and 1:1 line in (c) and (d). (a) Relationship between ln(chlorophyll a biomass) and principal component score based on <br />physical habitat parameters; (b) relationship between ln(invertebrate biomass) and principal component score based on physical <br />habitat parameters; (c) predicted and measured ln(chlorophyll a biomass), regression based on principle component score <br />and rk location; (d) predicted and measured ln(invertebrate biomass), regression based on principle component score and <br />ln(chlorophyll a biomass). <br />brate biomass and the physical variables that charac- <br />terize the riverbed (Table 2). Positive correlations were <br />found with variables reflecting less sedimentation <br />(DFS, D50, void volume) and higher detrital content <br />and water velocity, while inverse relationships were <br />found with variables reflecting higher sedimentation <br />(percentage of embeddedness and percentage of sub- <br />strate <2 mm). <br />The high degree of correlation among physical pa- <br />rameters (Table 2) made it difficult to determine which <br />parameters most related to standing crops of primary <br />producers and invertebrate consumers. Hence, PCA <br />factor scores were used to assess relationships between <br />physical habitat and measures of benthic biota. Factor <br />scores for the first principal component for each riffle <br />and run were calculated using the factor loadings in <br />Table 3 for each sample reach and regressed against <br />the corresponding values for chlorophyll a and biomass <br />of invertebrates. Biological parameters at each sample <br />reach were averaged over the three sample periods. The <br />analysis combined riffles and runs because physical <br />habitat differences were included in the factor score. <br />Highly significant relationships were found between <br />both chlorophyll a (F,,66 = 57.8, P < 0.0000001, r' = <br />0.47) and invertebrates (F,,66 = 50.3, P < 0.0000001, <br />rz = 0.43) and the principal component scores (Fig. 4a, <br />b). Regression,relationships were improved by adding <br />additional variables to the regressions containing the <br />factor scores. Because a highly significant (F,,66 = <br />130.2, P < 0.0000001, r2 = 0.66) relationship was also <br />found between chlorophyll a and invertebrate biomass <br />(Fig. 5a), and longitudinal relationships were found for <br />both chlorophyll a and invertebrates, addition of these <br />elements was evaluated. By including the distance up- <br />stream (rk) of the Colorado and Green River conflu- <br />ence, the regression relationship (Fig. 4c) for chloro- <br />phyll a improved (F2,59 = 104.6, P < 0.00001, rz = <br />0.78), and the multiple regression analysis indicated no <br />collinearity problems (condition number = 3.11; Bel- <br />sley et al. 1980). The relationship also improved for <br />invertebrates (Fig. 4d) when chlorophyll a was includ- <br />ed (F265 = 69.9, P < 0.00001, r2 = 0.68); however, <br />when rk location was added, the resulting condition <br />index of 17.4 suggested collinearity problems, proba- <br />bly reflecting the relationship between chlorophyll a <br />and rk location. <br />Algal invertebrate biomass-fish biomass relation- <br />ship.-Highly significant relationships were found be- <br />tween fish biomass (expressed as catch rates) and chlo- <br />rophyll a (Fig. 5b; F1,31 = 35.2, P < 0.00001, r' = <br />0.53), invertebrate biomass (F13, = 26.3, P = 0.00001, <br />r2 = 0.46), and combined chlorophyll a and inverte-