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<br />I <br /> <br />I <br /> <br />included extremely variable streamflows and high sediment loads. These fishes <br />evolved together and, more than likely, formed a stable fish community where <br />competition and predation were balanced. The main natural predator on the <br />native fishes in the Upper Colorado River Basin was the Colorado squawfish. <br />The other native species occupied various habitats (i.e., niches) that reduced <br />competition for food resources. <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />Today, the fish fauna in most areas of the Colorado River Basin bears little <br />resemblance to the fish community that occurred historically. Nonnative fish <br />species now compose 76% (42 of 55 species) of the fish community in the Upper <br />Colorado River Basin (Tyus et al. 1982). Since the nonnative fishes did not <br />evolve with the endangered fishes, mechanisms to balance predation and <br />competition to allow co-existence have not developed so nonnative fishes <br />predominate in the present fish community. This change in the fish community <br />is consistent with ecological theory where species that are more efficient in <br />capturing and converting food into biomass (i.e., nonnative species) will <br />persist over other species (i.e., native species) as resources become scarce <br />(Tilman 1982) . <br /> <br />I <br /> <br />I <br /> <br />III. ECOLOGICAL IMPORTANCE OF FLOODPLAINS <br /> <br />I <br /> <br />The importance of the land-water interface to the productivity of lotic <br />systems has been recognized for over twenty-five years (Allan 1995; Hynes <br />1970; Hynes 1983; Schlosser 1990, 1991; Ward 1989). However, interpretation <br />of the complexity of biological responses and importance of geomorphological <br />or hydrological processes has occurred only recently. <br /> <br />I <br /> <br />Historically, the ecological concepts of the river continuum (Vannote et al. <br />1980) and flood pulse (Junk et al. 1989) applied to the Upper Colorado River <br />Basin. The river continuum concept applied to the headwaters and high <br />gradient, restricted meander, canyon reaches while the flood pulse. concept <br />applied to low gradient, unrestricted reaches that form floodplains in broad <br />valleys. Lotic systems not only transfer organic matter from upstream reaches <br />in arid or semi-arid regions (i.e., continuum concept) but also deposit this <br />material in floodplains where these nutrients aid in high productivity of <br />invertebrates that periodically transfers food organisms to the main channel <br />of the river (i.e., flood pulse concept). Most of the productivity in a large <br />river-floodplain ecosystem occurs in the floodplain and is dependent upon the <br />duration of inundation and the area of inundation (Junk et al. 1989). With <br />dam construction, river ecosystems became fragmented (Ward and Stanford 1983, <br />1995) so that the continuum concept of energy transfer has been greatly <br />disrupted. Although the flood pulse concept was probably more important to <br />energy transfer than the continuum concept before Upper Basin rivers were <br />altered, the flood pulse concept is even more important today to productivity <br />of the present fragmented river ecosystem where extensive levees continue to <br />separate much of the floodplain habitat from main channels. The major zone of <br />productivity in a floodplain is the "moving littoral" (i.e., a shallow zone <br />that extends from the edge of the waterline to several meters in depth) <br />because it covers the maximum area of a floodplain for a given flood as it <br />traverses the floodplain during inundation and draining (Junk et al. 1989). <br /> <br />Floods and floodplains are now understood to be essential components of large <br />river systems (Bayley 1991; Petts and Maddock 1994; Sedell et al. 1989; <br />Welcomme 1995). The energy dynamics of large rivers is strongly influenced by <br />floodplain habitats (Sedell et al. 1989) where productivity is higher than in <br />river channel habitats (Hynes 1970; Welcomme 1985; Welcomme 1989). <br />Floodplains provide a greater opportunity for retention and processing of <br />nutrients and organic matter than main river channels (Sparks 1995). The <br />spawning strategies of fishes in many tropical and temperate areas are <br />correlated with the flood pulse that is associated with high productivity in <br />shallow, flooded areas (Copp 1989; Junk et al. 1989). Ephemeral wetlands in <br />arid regions (e.g., floodplains, arroyos, and playa lakes) are not included in <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br /> <br />I <br />I <br /> <br />9 <br />