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1942 <br />JOHNSON ET AL. <br />and Minckley 1991; Olden et al. 2006), and over 40 of <br />these are established in the upper basin alone. <br />Preventing introductions has proven to be very <br />difficult, and it is now recognized that coping with <br />invasions will require the ability to predict range <br />expansions and the intensity of effects based on life <br />histories of native and invasive species and character- <br />istics of recipient systems (Moyle and Light 1996; <br />Ricciardi and Rasmussen 1998; Kolar and Lodge 2001; <br />Iguchi et al. 2004; Vander Zanden et al. 2004; Bestgen <br />et al. 2006). Case studies of the relative effects of <br />invasions by multiple species can provide valuable <br />insights that will assist managers in anticipating and <br />controlling new invaders. <br />Debate continues regarding the relative importance <br />of habitat loss versus nonnative fishes in rivers of the <br />region, but the two factors usually operate concurrently <br />and even synergistically and are thus confounded. The <br />Yampa River is unique in that it allows for an <br />assessment of the magnitude of nonnative fish effects, <br />as it has the most natural hydrograph of any river of its <br />size in the upper Colorado River basin (Boehm 2004). <br />The lack of main-stem dams below the headwaters and <br />the absence of large diversions have maintained the <br />natural spring peak in discharge, although summer <br />flow has been reduced by agricultural withdrawals <br />(Stewart et al. 2005). Despite the availability of <br />relatively high-quality habitat, populations of small- <br />bodied native fishes and juvenile life stages of larger <br />species have declined precipitously during the past <br />decade. Native speckled dace Rhinichthys osculus, <br />mottled sculpin Cottus bairdii, bluehead suckers <br />Catostomus discobolus, and flannelmouth suckers <br />Catostomus latipinnis were strikingly lower in abun- <br />dance or were extirpated in some reaches of the Yampa <br />River, and nonnative fish abundance was higher in <br />2002-2003 than in 1998-1999 (Anderson 2005). <br />Predation by nonnative fishes has been implicated in <br />the decline of small-bodied fishes and may also be <br />responsible for local declines in large-bodied species, <br />such as Colorado pikeminnow (Bestgen et al. 2007). <br />However, information on the relative predatory impact <br />of various introduced species was lacking, and control <br />of myriad species would be impractical. Further, some <br />of these species are popular sport fish, so their control <br />would require explanation and justification to the <br />public. <br />One widely held opinion was that northern pike Esox <br />lucius posed the gravest threat to native fishes of the <br />Yampa River. This assumption presumably arose <br />because northern pike are notoriously piscivorous, <br />large, and able to prey on a wide range of fish sizes <br />(Scott and Grossman 1998). Diet studies in the system <br />confirmed that northem pike preyed upon native fishes <br />(Tyus and Beard 1990). Channel catfish Ictalurus <br />punctatus were also considered a threat, both as <br />predators and as competitors (Tyus and Nikirk 1990; <br />Tyus and Saunders 2000). Removal programs for both <br />species were implemented in 1998 (Hawkins et al. <br />2005; Mueller 2005). The potential impact of small- <br />mouth bass Micropterus dolomieu on native fishes may <br />have been discounted, because historically they were <br />very rare. Additionally, many sources (Carlander 1977; <br />Scott and Grossman 1998; Tomelleri and Eberle 1990; <br />Sigler and Sigler 1996) have noted the predominance <br />of crayfishes in the diet of smallmouth bass; this notion <br />is well entrenched in sporifishing lore. An unprece- <br />dented increase in the abundance of smallmouth bass in <br />parts of the Yampa River during the past decade <br />prompted interest in determining their diet and the <br />predation pressure they exerted on native fishes in the <br />system relative to predation by channel catfish and <br />northem pike. A predominately local but avid angling <br />clientele developed as abundance of smallmouth bass <br />increased, necessitating objective information and a <br />thorough analysis of the effects of this nonnative <br />predator before management actions aimed at reducing <br />its abundance could be considered. <br />Assessing the potential predatory impacts on native <br />species by channel catfish, northem pike, and small- <br />mouth bass requires information on the diets of each <br />predator species. However, to determine the relative <br />intensity of piscivory on prey populations, information <br />on consumption rates and demographics of each <br />population are also required. Further, in some situa- <br />tions, predation may have already depleted prey <br />species of concern and predators will have switched <br />to alternate prey; in those cases, contemporary diet <br />information gives a misleading impression of preda- <br />tors' potential impacts on species of concern. Bioen- <br />ergetics models can integrate ecological, physiological, <br />and demographic information to compute consumptive <br />demand of predator populations, a relevant metric <br />when assessing relative impacts. A great strength of <br />simulation modeling is that alternative scenarios can be <br />evaluated to ask questions such as, "What if native <br />prey fish populations rebound and predators begin <br />feeding on them again?" or "How many of those prey <br />could the predators consume?" <br />In this study, our objectives were to (1) assemble <br />available data on diets and demographics of three <br />nonnative predators from the Yampa River and diet <br />information from a reference system and (2) employ <br />bioenergetics models to quantify the relative predatory <br />threat of each species. The information was then <br />provided to managers so that predatory fish control <br />efforts could be prioritized and applied in an <br />ecologically and fiscally efficient manner and so that <br />