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28 BIOLOGICAL REPORT 24 <br />Life stages of target biota are sampled or otherwise <br />monitored (fish preferences are often determined <br />from animals fitted with radio transmitters) across <br />the range of the hydraulic variables to derive "habi- <br />tat suitability curves." Intuitively, this is a logical <br />approach, but it is often biased by sampling error, <br />especially in large, deep, and often turbid rivers, <br />where the biota are difficult to capture or see; (3) <br />the net suitability of use of a given locality (transect <br />cell) is quantified by a variable called weighted <br />usable area (WUA), which is a derived relation <br />between plan area of the transect cell (area avail- <br />able) and the habitat preference indices (from suit- <br />ability curves) for velocity, depth, and substratum. <br />The WUA is calculated cell by cell and summed for <br />the entire reach and over a range of discharges. <br />Hence, increments of WUA for a stream become a <br />continuous function of discharge. Easy to read and <br />more detailed descriptions of the IFIM are given by <br />Gore and Nestler (1988) and Nestler et al. (1989). <br />This procedure has been widely used to justify flow <br />provisions in regulated streams throughout North <br />America, in some cases leading to state statutes to <br />guarantee protection of aquatic biota (Reiser et al. <br />1989a). <br />Even though the IFIM has become an industry <br />standard (Reiser et al. 1989a), it has a number of <br />faults that are not widely recognized or under- <br />stood within management circles. Concern exists <br />regarding use of suitability curves as probability <br />functions (Patten 1979; Mathur et al. 1985; Moyle <br />and Baltz 1985); the assumption of independence <br />of depth, velocity, and substratum (Patten 1979; <br />Malthur et al. 1985); the lack of a demonstrated <br />relation between WUA and a meaningful measure <br />of productivity or biomass (Mathur et al. 1985; <br />Bowlby and Roff 1986; Conder and Annear 1987; <br />Scott and Shirvell 1987); and lack of any relation- <br />ship with regard to many other ecosystem proc- <br />esses, such as predation and other density-de- <br />pendent relationships, which clearly influence <br />population structure (Moyle and Baltz 1985; <br />Bowlby and Roff 1986; Orth 1987; Stanford and <br />Ward 1992a). To my knowledge none of these <br />criticisms has been resolved, nor is it likely they <br />will be. However, these criticisms have been <br />placed in perspective with respect to the rationale <br />and intent of the IFIM, which is often misunder- <br />stood, misrepresented, and misused (Gore and <br />Nestler 1988). For example, the model was not <br />intended to predict biomass; it is a physical habi- <br />tat simulator. Even when the model is ap- <br />plied properly, a variety of problems may emerge <br />depending on input choices, which necessitates a <br />clear understanding of how the model works. The <br />simulator can use a variety of hydraulic predictors <br />(e.g., the HEC-2 flow model of the U.S. Army <br />Corps of Engineers), each of which has biases and <br />therefore will result in different WTJA calculations <br />(Gan and McMahon 1990). Suitability curves not <br />derived on site (i.e., curves given in the literature) <br />are often used, which can also bias output (Gore <br />and Nestler 1988). <br />The IFIM was used in an attempt to derive flow <br />recommendations for specific river segments of <br />the Upper Colorado River Basin for the endan- <br />gered fishes. However, in the analysis WUA often <br />was maximized for various life history stages of <br />squawfish and humpback chub at very low flows <br />that in the historical record were exceeded most <br />or all of the time (Rose and Hann 1989). Such <br />output is nonsense because the ecological data for <br />these fishes clearly shows the importance of back- <br />waters and eddies that occur at much higher <br />flows. The problem here is that the IFIM probably <br />should never have been used in the big river <br />reaches of the Upper Colorado River Basin. When <br />low velocity habitats are abundant, as they are <br />throughout the potamon of the Colorado River <br />system, the simulator underestimates the WUA; <br />in fact, the model cannot deal with zero-flow habi- <br />tats. This explains why the IFIM works well only <br />in small streams where the channel is charac- <br />terized by uniformly varying flow (e.g., the low <br />velocity profile reflects steady, uniform flow, <br />which is also an assumption of the HEC-2 hydrol- <br />ogy simulator that is often used in IFIM; my <br />observations). Also, habitat suitability curves <br />were probably biased because the fish were diffi- <br />cult to observe or collect in the usually turbid, <br />deep water of the Yampa and Green rivers (Rose <br />and Hann 1989), which is precisely why the adult <br />fish monitoring program (U.S. Fish and Wildlife <br />Service 1987b) emphasizes shallow, shoreline <br />habitats that can be sampled effectively by elec- <br />trofishing. However, the fishes routinely use deep- <br />water habitats (e.g., Tyus and McAda 1984; <br />McAda and Kaeding 1991), and movement be- <br />tween habitats (e.g., channel, backwaters) on a <br />diel basis cannot be accounted for in the method. <br />The utility of the IFIM evolved a great deal during <br />the period that data were being gathered in the <br />Upper Colorado River Basin studies, and deficien- <br />cies in the method with regard to the Colorado <br />River were probably not apparent at the time <br />much of the data were gathered.