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6 BIOLOGICAL REPORT 24 <br />of nonnative fishes that may migrate upstream, <br />thereby influencing the native fishes living in the <br />Upper Colorado River Basin. However, I viewed <br />Lake Powell as the downstream boundary of the <br />river ecosystem examined in this report (Fig. 1). <br />A vital characteristic of river ecosystems is that <br />their biophysical processes are inherently vari- <br />able. The essence of ecology is understanding the <br />complex processes that control observed variabil- <br />ity in the distribution and abundance of biota. <br />Quantification of the structure and function of <br />complex systems, like the Upper Colorado River <br />Basin ecosystem, in time and space must be based <br />on long-term (> 5 years) measurements to detect <br />patterns or trends that in shorter time frames are <br />overwhelmed by variability. Hence, an ecosystem <br />approach strives to determine how and why the <br />river changes in time and space, not simply to <br />describe current conditions. <br />Like most scientists, I view model building and <br />logistic descriptions of dynamic events in ecology <br />as mechanistic tools for formalizing a better under- <br />standing of what is known about a system; such <br />tools should not be used to predict the future. <br />Predicting the consequences of environmental <br />change is the ultimate challenge of contemporary <br />ecology. This must be resolved through strong in- <br />ferences based on properly scaled measurements <br />of biophysical variables that integrate the myriad <br />system-specific ecological processes that are spa- <br />tially and temporally dynamic (Magnuson 1990; <br />Stanford and Ward 1992a). The problem of in- <br />stream flow provision must be resolved from <br />strong inferences derived from long-term trends in <br />ecological processes and responses of the river <br />ecosystem in which the endangered fishes live. <br />River Ecology and Effects of <br />Regulation on the Endangered <br />Fishes of the Upper Colorado <br />River Basin <br />Ecology of the Endangered Fishes <br />Information about the endangered fishes is very <br />detailed, given that they are relatively rare fishes; <br />several reviews of the scientific information have <br />been published (e.g., Stanford and Ward 1986b; <br />Minckley et al. 1991; Tyus 1991a). Therefore, I <br />repeat here only salient points of particular impor- <br />tance to my review of the flow recommendations <br />made by the U.S. Fish and Wildlife Service. <br />As noted above, the historical range of the four <br />species included the potamon and transitional <br />reaches of the Green and Colorado river systems, <br />including most of the larger tributaries, in particu- <br />lar the Yampa, White, Dolores, and Gunnison riv- <br />ers. Today, ranges of these fish are fragmented by <br />dams and diversions, and populations have de- <br />clined significantly in relation to distributions at <br />the turn of the century (Quartarone 1993). <br />Bonytail chub are close to extirpation, but they <br />have been successfully cultured, along with hump- <br />back chub, squawfish, and razorback sucker, at the <br />Dexter National Fish Hatchery, Dexter, New Mex- <br />ico (Johnson and Jensen 1991), and brood stocks <br />currently are being held in several locations. Be- <br />cause of their comparative rarity in the wild, eco- <br />logical information on the historical range of <br />bonytail chub is more fragmentary than for the <br />other species. A few specimens of bonytail chub <br />were collected in the 1970's in the Green and <br />Yampa rivers (Kaeding et al. 1986), but their phe- <br />nology (life history) and exact cause of disappear- <br />ance in the Upper Colorado River Basin system are <br />unknown. <br />Humpback chub are found only in whitewater <br />canyon segments (Fig. 1). Migrations are limited, <br />and humpback chub may have always been re- <br />stricted to specific canyon segments, at least as <br />adults. Spawning in the Upper Colorado River Ba- <br />sin occurs on the declining limb of the spring runoff <br />event in association with the 20° C isotherm <br />(Kaeding and Zimmerman 1983). Humpback chub <br />interact behaviorally (and probably hybridize) with <br />congeneric, endemic roundtail chub (Gila robusta), <br />which are more abundant throughout the Upper <br />Colorado River Basin (Kaeding et al. 1990; Karp <br />and Tyus 1990). Much of what is known about the <br />life cycle of humpback chub is based on unpublished <br />studies in the Grand Canyon, where they migrate <br />from the regulated Colorado River into the unregu- <br />lated Little Colorado River to spawn. Similar mi- <br />gratory behavior has not been documented in the <br />Upper Colorado River Basin, and exact locations of <br />spawning sites are unknown (Richard Valdez, <br />BioWest Inc., Logan, Utah, personal communica- <br />tion; Larry Crist, U.S. Bureau of Reclamation, Salt <br />Lake City, Utah, personal communication). <br />Lanigan and Tyus (1989) estimated that only <br />978 ± 232 adult razorback sucker remained in the <br />Green River above Desolation Canyon during <br />1981-86, which very likely is only a small fraction <br />of the historic population. Some researchers be- <br />lieve that significant declines have occurred since