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December 2002 FLOW-SEDIMENT EFFECTS ON RIVERINE FISH <br />proportion of captured individuals ?550 mm TL, and <br />correction factors based on stratum-specific catch-per- <br />effort values and kilometers per stratum. Because of <br />the rarity of this species, all available data (1991-1994 <br />and 1998-1999) were used to characterize population <br />distribution including years outside of the 1994-1995 <br />study period. <br />Comparisons of fish body condition among strata <br />also were used as a means to assess food availability. <br />Only the three dominant large-bodied native species <br />were analyzed: Catostomus discobolus Cope (bluehead <br />sucker), C. latipinnis Baird and Girard (flannelmouth <br />sucker), and Gila robusta Baird and -Gerard (roundtail <br />chub). Relative condition (K) is the observed mass of <br />a given fish divided by the expected mass for a fish of <br />its length (Le Cren 1951). The expected mass is cal- <br />culated using constants (slope and intercept coeffi- <br />cients) derived from regressing log-transformed mass <br />as a function of length (see Osmundson et al. 1998). <br />We used length and mass of all captured individuals <br />from the last three sample periods to develop one <br />length-mass relationship for each species that could <br />then be used as a standard for among-strata compari- <br />sons. <br />Statistical procedures <br />Various statistical procedures were used including <br />analysis of variance (ANOVA), analysis of covariance <br />(ANCOVA), Pearson correlation, multiple regression, <br />and principal component analysis (PCA). All statistical <br />analyses were performed using NCSS (2000). In the <br />results, the test used for each analysis is stated along <br />with the pertinent statistics and P values. For statistical <br />analyses, log transformations (ln[x + 1]) were used to <br />increase- normality in biological parameters; all anal- <br />yses of chlorophyll a, macroinvertebrate biomass, and <br />fish numbers and biomass are from In-transformed val- <br />ues. For ANOVA tests, the Tukey-Kramer multiple <br />comparison test was used for post hoc testing among <br />treatments (P < 0.05). <br />PCA was used to explore relationships among phys- <br />ical attributes of the riverbed. The PCA attempts to <br />maintain the information of several variables with few- <br />er independent variables (often one to three). Values <br />(scores) of this reduced'set of variables are calculated <br />from transformed original data using loading coeffi- <br />cients (maximum absolute value = 1.0). Ranges in the <br />scores of the resulting variables are useful in charac- <br />terizing the original data set and relating to other var- <br />iables. We used PCA to characterize sites relative to <br />bed-sediment characteristics. The scores from the first <br />principal component from seven physical variables <br />were regressed against chlorophyll a and macroinver- <br />tebrate biomass to assess relationships between the <br />physical environment and the lowest trophic levels. The <br />seven physical variables were: fraction of the substrate <br /><2 mm, median particle size of the surface layer, vol- <br />ume of interstitial void space, fraction of the surface <br />1725 <br />area consisting of fines, total DFS, midcolumn water <br />velocity, and CPOM dry mass. <br />RESULTS <br />Longitudinal variation <br />Physical characteristics.-River gradient increased <br />almost exponentially in an upstream direction, varying <br />from 0.028% in stratum 1 to 0.196% in stratum 11 (Fig. <br />1, lower). The overall mean gradient of 0.100% was <br />equaled or exceeded in strata 7-11 and stratum 4. Bank- <br />full depth of the river channel increased systematically <br />downstream (Fig. 2a), whereas bankfull width varied <br />greatly among strata but with no longitudinal trend. <br />Mean bankfull depth doubled between strata 11 and 5 <br />and reached a maximum in canyon-bound stratum 3; <br />it then decreased through strata 2 and 1. Actual water <br />depth at base flow was not measured. Habitat mapping <br />indicated a steady, nearly linear, downstream decrease <br />in surface area of riffles, varying from 12.9% of total <br />surface area in stratum 11 to <0.1 % in stratum 1 (Fig. <br />2b). <br />Estimates of discharges necessary to initiate motion <br />of coarse bed materials (characterized by sporadic mo- <br />tion of a few particles somewhere on the bed) at 50% <br />of the cross sections increased with distance down- <br />stream, but were similar among strata bounded by the <br />same major tributaries (Table 1; Fig. 1); in strata 2- <br />11, these discharges averaged 40% of the bankfull dis- <br />charge (Pitlick and Cress 2000). Discharges necessary <br />to cause widespread bed mobilization at 50% of the <br />cross sections also increased downstream (-Table 1); in <br />strata 6-11, these discharges were similar among strata <br />bounded by the same major tributaries and correspond- <br />ed to the bankfull discharge. Downstream of the Do- <br />lores River inflow (strata 1-5), some very high dis- <br />charges are required to reach bankfull owing to large <br />increases in either channel width or depth, and thresh- <br />olds for widespread bed mobilization were assumed to <br />occur at discharges less than bankfull. <br />Other physical habitat characteristics were estimated <br />separately for riffle and run habitats. However, char- <br />acterization of riffles was precluded in stratum 1 be- <br />cause of the near absence of this habitat type. For the <br />whole study area, the mean Dso (median grain size) of <br />riffle substrates was greater than that of run substrates <br />(ANOVA, F,,gg = 21.12, P = 0.00001). Also, there was <br />a fairly continuous and significant (ANCOVA, F,,,, = <br />52.19, P < 0.00001) downstream decrease in the Dso <br />of runs, though not of riffles (ANCOVA, F1,72 = 0.96, <br />P = 0.33). In riffles, the DSO increased in the three <br />strata immediately downstream of stratum 6 (Fig. 2c). <br />In contrast, when fine sediments were excluded from <br />the analysis and only coarse substrate particles (>_2 mm <br />diameter) were considered (Fig. 2d), mean grain sizes <br />of riffle and run substrates were not significantly dif- <br />ferent (ANOVA, F,,,, = 0.50, P = 0.48), and mean <br />grain sizes in runs did not decrease with distance down-