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<br />recently. The concepts of the river continuum (Vannote et al. 1980) and flood <br />pulse (Junk et al. 1989) apply to the Upper Colorado River Basin. The river <br />continuum concept applies to the headwaters and high gradient, restricted meander <br />canyon reaches while the flood pulse concept applies to low gradient, <br />unrestricted reaches that form floodplains in broad valley reaches. Lotic <br />systems not only transfer organic matter from upstream reaches in arid or semi- <br />arid regions (i.e., continuum concept) but also deposit this material in <br />floodplains where high productivity of invertebrates periodically enters the <br />river (i .e., flood pulse concept). Shallow floodplain habitats become much <br />warmer than the adjacent river, increasing the productivity for phytoplankton and <br />development of a food web (Welcomme 1979). Slow growth and higher mortality of <br />endangered Colorado River fishes has been attributed to lower water temperatures <br />in the Upper Colorado River Basin (Kaeding and Osmundson 1988). Floods and <br />fl oodp 1 a ins are now understood to be essent i a 1 components of ri ver systems <br />(Bayley 1991; Petts and Maddock 1994; Sedell et al. 1989). The energy dynamics <br />of large rivers is strongly influenced by floodplain habitats (Sedell et al. <br />1989) where productivity is higher than habitats in river channels (Hynes 1970; <br />Welcomme 1985; Welcomme 1989). The spawning strategies of fishes in many <br />tropical and some temperate areas are correlated with the flood pulse that is <br />associated with high productivity in shallow, flooded areas where organic matter <br />is retained (Junk et al. 1989). Welcomme (1985) stated that the shallow littoral <br />zone of floodplain habitats produce higher densities of zooplankton when compared <br />with the entire floodplain area. <br /> <br />IV. RELATION OF NUTRIENTS, SUNLIGHT PENETRATION, AND <br />WARM WATER TEMPERATURES IN PHYTOPLANKTON PRODUCTION <br /> <br />Phytoplankton productivity provides the basis for development of a food web. <br />Phytoplankton production and standing crops increase in concert with increases <br />in annual input of nutrients regardless of latitude. Carbon, nitrogen, and <br />phosphorus are key elements for phytoplankton production. Phosphorus is the most <br />limiting element in north temperate and subarctic waters (Schindler 1978). <br />Nitrogen is the most abundant element in the atmosphere and is generally not <br />limiting. Also, abundant carbon dioxide in the atmosphere provides the necessary <br />carbon. Therefore, phytoplankton production and standing crop in north temperate <br />freshwaters is generally proportional to the phosphorus input. Particulate <br />phosphorus, either chemically desorbed or actively mobilized by microbiota, is <br />not readily available in rivers with a high sediment load because most of the <br />phosphorus is bound to the sediments (Ell is and Stanford 1988). Watts and <br />Lamarra (1983) determined that between 21% and 49% of the total phosphorus in <br />Colorado River water at the bridge upstream from Moab, Utah in September and <br />October 1978 was bioavailable with most of the extractable element in the form <br />of calcium-bound phosphorus. Therefore, Watts and Lamarra (1983) concluded that <br />algae production was not nutrient limited in this reach of the Colorado River but <br />that primary production in this reach was inversely related to the turbidity of <br />the riverine environment. <br /> <br />Turbidity from suspended fine sediments in Upper Colorado River Basin rivers is <br />high and adversely affects primary and secondary production. Product i on of <br />phytoplankton and zooplankton (Tables 1 and 2) that form the basis for a food <br />pyramid are extremely low in the these rivers (Grabowski and Hiebert 1989; Cooper <br />and Severn 1994 a, b, c, and d; Mabey and Schiozawa 1993). High turbidity in the <br /> <br />4 <br />