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1720 D. B. OSMUNDSON ET AL. Ecological Applications <br />Vol. 12, No. 6 <br />Fausch et al. 1988), yet many studies suggest that biotic <br />factors such as food availability may be more important <br />(Cada et al. 1987, Filbert and Hawkins 1995, Hughes <br />1998, Nislow et al. 1999). With the exception of the <br />flood-pulse concept, wherein main-channel productiv- <br />ity is enhanced by the entrainment of floodplain nu- <br />trients and carbon (Junk et al. 1989, Bayley 1995, Pow- <br />er et al. 1995), few investigations have attempted to <br />link flow effects to availability of food for fishes. <br />To restore native fish populations in regulated rivers, <br />management strategies must be based on an under- <br />standing, not only of the fishes' life-history attributes, <br />but also of the ecosystem that sustains them (Power et <br />al. 1995, Stanford et al. 1996, Wootton et al. 1996). <br />Petts (1991) identified the need for integration of in- <br />terdisciplinary research that approaches the study of <br />regulated rivers on a longitudinal basis and in a hier- <br />archical manner: hydrology and geomorphology to de- <br />scribe the influence of flow regime and sediment sourc- <br />es on river structure, and ecology to link these influ- <br />ences to riverine fauna. However, large-scale riverine <br />processes are inherently difficult to study (Schlosser <br />1991, Johnson et al. 1995, Peterson and Kwak 1999). <br />With some exceptions, experimental studies are gen- <br />erally not possible (Power et al. 1995, Poff et al. 1997), <br />so correlative empirical approaches are required to dis- <br />cern linkages among flow, sediment, and biota. <br />The Colorado River, in southwestern United States, <br />is one of the most regulated rivers in North America <br />(Fradkin 1981). Nearly all of the pathways by which <br />regulation can alter or degrade native aquatic fauna are <br />represented, supplying diverse challenges for the ap- <br />plied ecologist. Driven by the mandates of the Endan- <br />gered Species Act, resource managers need solutions <br />to ecologically complex problems (Stanford 1994, <br />Holling and Meffe 1996). Herein, we report the results <br />of our investigations concerning, physical habitat and <br />trophic relations in a contiguous portion of the upper <br />Colorado River inhabited by all life stages of Ptych- <br />ocheilus lucius Girard (Colorado pikeminnow), an en- <br />dangered piscivorous cyprinid. For this population, low <br />recruitment and low adult carrying capacity are two <br />primary constraints that limit population viability (Os- <br />mundson and Burnham 1998, Osmundson et al. 1998). <br />In addition to the obvious effects of reduced range from <br />instream barriers, we hypothesized that the capacity of <br />the system to support adult pikeminnow may have de- <br />clined from a reduction in food availability due to in- <br />creased persistence of fine sediment (particles <2 mm), <br />a possible effect of river regulation. <br />We used an interdisciplinary approach to investi- <br />gating the role of flow regime in riverine trophic struc- <br />ture, integrating geomorphology with studies of fish <br />community structure and biomass of lower trophic lev- <br />els (Petts 1991, Johnson et al. 1995). We first described <br />the distribution of physical habitat and biota on a lon- <br />gitudinal basis throughout the present and historical <br />range of Colorado pikeminnow in the upper Colorado <br />River mainstem. Then, using empirical relations, we <br />sought links among sediment distribution, standing <br />crops of primary producers, consumers, and the prin- <br />cipal piscivore (Colorado pikeminnow). The relation <br />between flow and sediment movement was next as- <br />sessed through standard geomorphological analyses. <br />Utilizing these empirical relations and historical flow <br />records, we evaluated the potential for reduced predator <br />food availability as affected by functional changes in <br />flow. Our goal was to assess links among flow, sedi- <br />ment, and biota in a large river system and determine <br />if such linkages might provide a food-mediated path- <br />way by which flow regime alteration affects fish as- <br />semblage structure. We discuss the implications for na- <br />tive fish restoration efforts in general and provide man- <br />agement recommendations for conservation of Colo- <br />rado pikeminnow in the upper Colorado River. <br />SPECIES OVERVIEW AND STUDY AREA <br />Colorado pikeminnow <br />The Colorado pikeminnow, a large, long-lived pis- <br />civore, along with three other large-river, warm-water <br />fish species (two chubs, Gila cypha Miller and G. ele- <br />gans Baird and Girard, and one sucker Xyrauchen tex- <br />anus Abbott) endemic to the Colorado River Basin, are <br />classified as endangered under the Endangered Species <br />Act (U.S. Fish and Wildlife Service 2000). All four <br />once occurred throughout the basin, ranging from Wy- <br />oming south to the Gulf of California (Miller 1961). <br />Due primarily to extensive dam construction during the <br />1930s to 1960s, populations of Colorado pikeminnow <br />were extirpated in the lower sub-basin (downstream of <br />Lee's Ferry) by the 1970s and today occur only up- <br />stream of Glen Canyon Dam, primarily in the states of <br />Utah and Colorado (Minckley and Deacon 1968, <br />Minckley 1973). In the upper sub-basin, the range of <br />this species was reduced early in the century by di- <br />versions built for local irrigation projects, and addi- <br />tionally by the more recent (1960s-1980s) construction <br />of several large dams. Lotic environments were con- <br />verted to lentic habitats, downstream water tempera- <br />tures were reduced by hypolimnetic releases, and mi- <br />gration routes were blocked (Ono et al. 1983). After a <br />75% basin-wide loss of range, a viable population oc- <br />curs today only in the Green River system (Gilpin <br />1993). A smaller, less viable population occurs in the <br />mainstem Colorado River and its Gunnison River trib- <br />utary (Osmundson and Burnham 1998). Populations of <br />Ptychocheilus lucius appear to require an extensive <br />length of river with an array of habitat types to meet <br />the changing needs of different life stages. Larvae <br />hatched in gravel-cobble substrates of high-gradient <br />reaches drift 100-200 km downstream to low-gradient <br />reaches where backwaters formed in silt-sand bars pro- <br />vide ideal nursery habitat (Haynes et al. 1984, Tyus <br />and Haines 1991). Insectivory is largely replaced by <br />piscivory during the first year (Vanicek and Kramer