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24 BIOLOGICAL REPORT 24 <br />new rhithron food web, including the valuable <br />trout fishery, was severely damaged by the epi- <br />sodic side flows that occurred during summer <br />1991 and 1992, when the regulated flows were at <br />or near the 300 cfs minimum. Benthos and fish <br />were smothered by fine sediments, a situation <br />that has persisted owing to the lack of a spring <br />flush to clean the substratum (my observation). <br />Recent experimental flows to help determine flow <br />recommendations for endangered fishes in the <br />Gunnison River reached 4,000 cfs in 1992 but <br />were insufficient to rearrange alluvium entrained <br />in the river channel (Elliott and Parker 1992). <br />Because of the interactive effects of (1) a lack of <br />spring peaks or other flushing flows, (2) an ex- <br />tended period of minimum flow (both 1 and 2 due <br />to drought and regulation), (3) warmer tempera- <br />tures associated with low flows, and (4) episodic <br />loading of the channel from ephemeral side flows, <br />the position of the discontinuity moved upstream <br />during 1991-92, and side channels and eddies <br />filled with fine sediments and vegetation. Today, <br />the riparian corridor of the river is densely vege- <br />tated, and surface water and groundwater ex- <br />change with critically important backwater sys- <br />tems (e.g., Fig. 2) has been altered or lost <br />(Stanford and Ward 1992b). The food web in the <br />lower part of the Gunnison Gorge remains im- <br />paired owing to persistent fine sediments in and <br />on the substratum, which prevents establishment <br />of a productive biofilm and restricts attachment <br />sites for zoobenthos. <br />The Gunnison River case history illustrates a <br />classic response of a stream to regulation. Similar <br />results have been recorded elsewhere (e.g., Petts <br />1986; Stanford and Hauer 1992). An upstream <br />discontinuity exists on the Colorado River (Voelz <br />and Ward 1991) and the Green River (Pearson and <br />Franklin 1968; Pearson et al. 1968), although the <br />latter is significantly reset toward predam pota- <br />mon conditions by the Yampa River (Annear and <br />Neuhold 1983). <br />Conclusions Based on Review of the <br />Ecological Literature Pertaining to the <br />Endangered Fishes and the Regulation <br />of Flow <br />The endangered fishes remain relatively rare <br />in the Upper Colorado River Basin as a conse- <br />quence of stream regulation and possibly pre- <br />dation and other interactions with nonnative <br />fishes. Recruitment of adults has not been <br />clearly demonstrated for any of the species, but <br />age structure of squawfish suggests adult re- <br />cruitment is occurring (i.e., larvae, YOY, juve- <br />niles, and adults are collected each year in the <br />Upper Colorado River Basin, although all age <br />classes often are not observed in the same river <br />segments). Clear evidence for adult recruit- <br />ment is lacking for the other species. In recent <br />years gravid razorback sucker and humpback <br />chub were collected during the spawning sea- <br />son at a few sites, and a few YOY were col- <br />lected. Bonytail chub seem to be extirpated. <br />2. The distribution, relative abundance, and some <br />important physical habitat preferences of <br />squawfish, humpback chub, and razorback <br />sucker (in that order) are reasonably well <br />known (Fig. 6) and documented in peer-re- <br />viewed literature. However, only the life his- <br />tory of squawfish is fairly well understood. <br />Important aspects of the life history and habi- <br />tat preferences for humpback chub and razor- <br />back sucker remain to be documented. Much of <br />what is known about the life history and popu- <br />lation dynamics of humpback chub is based on <br />unpublished studies in the Grand Canyon, <br />which may or may not be relevant to the Upper <br />Basin (e.g., no population in the Upper Basin <br />is known to migrate into a tributary to spawn, <br />as occurs in the Little Colorado River within <br />the Grand Canyon; Larry Crist, personal com- <br />munication). Detailed information about <br />spawning and rearing is lacking for humpback <br />chub and razorback sucker throughout the Up- <br />per Basin, and virtually nothing is known <br />about bonytail chub. Moreover, accurate esti- <br />mates of annual population size are problem- <br />atic for all of the fishes (Tyus 1992), and mark- <br />recapture studies using the new transponder <br />tag technology are warranted. On the other <br />hand, a great deal more is known about the <br />distribution and abundance of the fishes, ex- <br />cept bonytail chub, than is known about the <br />influences of river hydraulics, sediment trans- <br />port, and riparian controls on the food web that <br />supports the fishes. In other words, the data on <br />which current flow recommendations are <br />based primarily describe the distribution and <br />abundance of the fishes, not the ecosystem- <br />level processes and responses that determine <br />productivity. <br />3. Strong linkages between trophic (food web) and <br />geomorphic attributes of the Upper Colorado <br />River Basin ecosystem are variable in time and <br />space. For example, algae (periphyton) and