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BIOLOGY OF HUMPBACK CHUBS IN THE GRAND CANYON <br />ruary, 74% of 57 in April, and 17% of 23 in <br />May. Nuptial tubercles were observed on 50, <br />23, and 22% of the fish in these respective sam- <br />ples. <br />Observations on growth rate of humpback <br />chubs spawned and raised in a hatchery (Ham- <br />man 1982) indicated that fish 14-18 mm long <br />from the Little Colorado River part of the con- <br />fluence (C 4) in May 1981 (Fig. 3) probably <br />resulted from spawning 2 or 3 weeks earlier. <br />Humpback chubs from the Little Colorado, <br />Colorado, and the confluence exhibited post- <br />ovulatory characteristics of low gonadosomatic <br />indices in May, and some large residual ova. <br />Gonad-development stage appeared similar in <br />histological preparations from these concurrent <br />collections (Charlie Smitb, Fish Cultural De- <br />velopment Center, USFWS, Bozeman, Mon- <br />tana, personal communication). These data sug- <br />gest that female humpback chubs undergo <br />seasonal gonad development in the Colorado <br />and may spawn there. This observation was not <br />unexpected. Temperatures of the Colorado <br />during the period of rapid seasonal gonad de- <br />velopment are similar to those present before <br />Glen Canyon Dam was constructed (Fig. 2). We <br />found one male humpback chub with nuptial <br />tubercles and running milt among eight hump- <br />back chubs collected in an overnight set of a <br />trammel net in a Colorado River (C 3) back- <br />water in May 1981. <br />Although humpback chubs might spawn in <br />the Colorado, our data strongly suggest that <br />spawning there does not result in the produc- <br />tion of viable offspring and the recruitment of <br />young fish to the population. No humpback <br />chubs shorter than 145 mm were collected from <br />the Colorado upstream from the confluence, <br />even though mature fish were present in this <br />river reach (Fig. 3). This distribution suggests <br />that small humpback chubs in the Colorado re- <br />sulted from spawning in the Little Colorado. <br />This hypothesis is supported by results of recent <br />thermal tolerance tests that showed tempera- <br />tureslike those of the Colorado River study area <br />preclude appreciable reproduction of hump- <br />back chubs. Hamman (1982) found that at 12- <br />13 C, only 12% of fertilized humpback chub <br />eggs hatched after 340-475 hours of incubation <br />and only 15% of these reached the feeding-larva <br />stage in a controlled hatchery environment. <br />When water temperature was maintained at 16- <br />17 C, incubation time was about 167-266 hours; <br />585 <br />TABLE 2.-Food arga~nisrns in the stomachs of humpba.ck~ <br />chubs collected from the Little Colorado and Colorado <br />rivers. Data are mean percentages of the total number of <br />organisms per sample, and mean frequencies (percent of <br />stomachs) within saraptes for each food organism. A sam- <br />ple is the group of stomachs (including those that ruere <br />empty) collected during a season (quarter year). Ranges <br />for samples are in parentheses. T =trace (<0.5%). <br />Little <br />Colorado Colorado <br />Taxon or (4 samples; (2 samples; <br />measure 26 fish) 18 fish) <br />Number of organisms <br />per stomach 7 (3-14) 192 (49-336) <br />Chironomidae <br />% numbers 36 (6-92) 28 (T-55) <br />%frequency 37 (20-50) 50 (1 I-89) <br />Simuliidae <br />% numbers 22 (0-90) 71 (43-99) <br />% frequency 5 (0-20) 56 (33-78) <br />Other Dipterae <br />% numbers 12 (0-36) T (T) <br />%frequency 22 (0-50) 11 (Ll) <br />Trichoptera <br />% numbers 18 (0-67) T (T) <br />% frequency 16 (0-50) 6 (0-1 ] ) <br />Neuroptera <br />% numbers 2 (0-10) 0 <br />% frequency 7 (0-27) 0 <br />Coleoptera <br />% numbers 2 (0-6) T (T) <br />%frequency 18(0-50) 11(11) <br />Gammarus <br />% numbers 0 1 (T-2) <br />%frequency 0 22 (I1-33) <br />Other° <br />% numbers 7 (0-17) T (T) <br />frequency 35 (0-100) 11 (11) <br />a Geratopogonidae, Hernerodromia, Ephydridae, Limno- <br />phora. <br />n Ephemeroptera, Orthoptera, Hymenoptera, Oligochae- <br />ta, Nematoda, Pirnephales firoraelas. <br />hatching success was 62%, and 91% of the em- <br />bryos that hatched became feeding larvae. <br />Humpback chub reproduction in the Colorado <br />is made more improbable by the frequent water- <br />level fluctuations (Fig. 2). A decline in water <br />level would expose fertilized eggs deposited in <br />shallow lentic areas-the only locations where <br />the water might be warm enough to otherwise <br />allow reproduction. <br />Stomach Contents <br />Stomach contents from humpback chubs were <br />numerically dominated by immature Chiro- <br />