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INSTREAM FLOWS TO ASSIST THE RECOVERY OF ENDANGERED FISHES 21 <br />windows (Fig. 14) determined to maximize areas <br />of backwater habitats in the alluvial nursery ar- <br />eas of the Green River during summer and fall <br />1992, peaking operations still caused considerable <br />diel fluctuation of river stage (e.g., Figs. 12-14). I <br />infer that backwaters thought to be protected by <br />these flow windows were in fact flushed or, at <br />least, significantly fluctuated repeatedly during <br />late summer 1992. Data presented by Grabowski <br />and Hiebert (1989) indicate that the food webs in <br />the backwater environments of the Green River <br />are not very productive. As noted above, these <br />backwaters should contain rooted aquatic plants <br />and a biodiverse, productive invertebrate and fish <br />food web. I realize that the fluctuations shown in <br />Figs. 12-14 are considerably reduced from opera- <br />tions in the past. Nonetheless, development of <br />stable, productive food webs in the backwaters <br />probably has not occurred as a consequence of <br />reregulation of the Flaming Gorge releases. More- <br />over, these backwaters probably will never be very <br />productive unless flow fluctuations can be elimi- <br />nated. Empirical information with which to firmly <br />judge the productivity of backwater food webs as <br />influenced by regulated baseflow regimes <br />throughout the Upper Colorado River Basin <br />is sorely needed and should be approached in <br />the dynamic time and space context described <br />above. <br />Peaking operations at Flaming Gorge are at- <br />tenuated in relation to distance downstream from <br />the dam. Therefore, baseflow instability (Figs. 12 <br />and 13) progressively worsens upstream from Jen- <br />sen and may be severe in the Echo and Brown Park <br />reaches. Elsewhere between Jensen and the dam, <br />the river is constrained in canyons, and the problem <br />may be somewhat ameliorated by geomorphology. <br />However, peaking flows are known to interrupt <br />insect emergences that feed the trout fishery in Red <br />Canyon immediately downstream from the dam <br />(my observation and Larry Crist, personal commu- <br />nication). Similar effects were observed on the Mis- <br />souri River below Holter Dam in Montana, and an <br />outcry from fly fishermen caused load control op- <br />erations to be shifted to another dam. The effect <br />was a translocation of stream regulation effects <br />from one river to another, thereby confounding <br />management objectives (Stanford and Hauer <br />1992). This illustrates the potential difficulty of <br />changing dam operations to meet the needs of en- <br />dangered fishes in potamon reaches of the Upper <br />Colorado River Basin, if rhithron trout fisheries <br />might be influenced in the process. <br />Stream Regulation Mediates Invasions of <br />Nonnative Predators and Complicates <br />Provision of Instream Flows to Protect <br />Endangered Fishec <br />Introduction of trout and other nonnative fish <br />in regulated streams is an enormously confound- <br />ing problem in the interpretation of the ecology of <br />regulated streams because the native species vir- <br />tually always seem to decline in the presence of <br />exotics, especially if the river is regulated. This <br />pervasive ecological problem has been reviewed <br />thoroughly (e.g., Mooney and Drake 1986). Preda- <br />tion of natives, including endangered fishes, by <br />exotics does occur in the Upper Colorado River <br />Basin, and red shiner, fathead minnow (Pi= <br />mephales promelas), walleye (Stizostedion <br />vitreum), northern pike (Esox lucius Linnaeus), <br />channel catfish, largemouth bass (Micropterus <br />salmoides), smallmouth bass (Micropterus <br />dolomieui), and green sunfish are especially prob- <br />lematic invaders (cf., Karp and Tyus 1990; Tyus <br />1991b; Tyus and Haines 1991). However, Meffe <br />(1984) and Minckley and Meffe (1987) showed <br />that intense flooding in rivers in the southwestern <br />United States was positively correlated with di- <br />versity and abundance of native fishes and nega- <br />tively correlated with diversity and abundance of <br />nonnative fishes. The strong inference is that non- <br />natives are maladapted to survive intense and <br />frequent (annual, at least) flooding compared with <br />natives. Having fewer predators increases re- <br />cruitment of natives and over time allows the <br />natives to persist in greater abundance than non- <br />natives (Fig. 15). The work of Meffe and Minckley <br />included the Virgin River and other tributaries of <br />the Colorado River but none in the upper basin. <br />Thus, while the data are not directly applicable, <br />the relationship probably holds. Hawkins and <br />Nesler (1991) correlated lower ratios of nonna- <br />tives to natives with high peak flows in the Yampa <br />River, and red shiner populations declined after <br />years of high spring flows in the Colorado River <br />(Osmundson and Kaeding 1991). <br />The prediction that flooding will limit preda- <br />tion mortality of endangered fishes is used as one <br />rationale in the recovery program for reinstate- <br />ment of peak flows. However, introduced species, <br />red shiner for example, are native in rivers that <br />experience floods (of bankfull or greater) rather <br />frequently, which suggests that flow augmenta- <br />tion might not work very well in controlling some <br />nonnative species. However, the complex interac- <br />tions described above that are associated with