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-
|